
Glass Q /... QfJ 
Rnnk Sj"f~ 






ATTTHOPi'S EDITION. 



DEPARTMENT OF THE INTERIOR. 

UNITED STATES GEOLOGICAL AND GEOGRAPHICAL SURVEY 
F. V. HAYDEN, U. S. Geologist-in-Charge. 



C O N T R I B U T IONS 



TO TIIK 



VNATOMY OF BIRDS. 




BY 



R. W. SHUPELDT, M. IX, 

Captain, Medical Department, U. S. A., member of the Philosophical, Anthropological. 

and Biological Societies of Washington, Honorary Curator of the 

Section of Avian Osteology of the Smithsonian Institution. 



EXTRACTED FKOM THE TWELFTH ANXFAL REPORT OF THE SURVEY. 



Washington. October 14, V&&1. 



/ 



AUTHOR'S EDITION. 



DEPARTMENT OF THE INTERIOR. 

UNITED STATES GEOLOGICAL AND GEOGRAPHICAL SURVEY. 
F. V; HAYDEN, U. S. Geologist-in-Charge. 

_ <Pf2~ 

C O N T R I B U T IONS 



TO TI1K 



ANATOMY OF BIRDS. 




BY 

K. W. SHUFELDT, M. D., 

n 
Captain, Medical Department, U. S. A., member of the Philosophical, Anthropological , 
and Biological Societies of Washington, Honorary Curator of the 
Section of Avian Osteology of the Smithsonian Institution. 

EXTRACTED FROM THE TWELFTH ANNUAL REPORT OF THE SURVEY. 



Washington, October 14, 188:J. 






G^* 



*4%» 



0) 



OSTEOLOGY OF SPEOTYTO CUNICULARIA HYPOGJIA. 



By R. W. Shufeldt, M. D., 

Captain, Medical Department, United States Army. 



At the present writing we know of but one species of the so-called 
Burrowing Owls inhabiting America, and this is represented by three 
existing races, the typical and largest of these being the Athene cunicu- 
laria, a species confined to South America, while its two varieties occur 
within the limits of 



the United States. 
Little difference 
seems to exist be- 
tween these latter, 
S. cunicularia flori- 
dana being of near- 
ly the same size as 
the subject of this 
paper, but darker 
in plumage, with 
fewer feathers 
upon the tarsi, so 
that, with the ex- 
ception of certain 
measurements, the 
description of the 

Skeleton Of any One Speotyto cunicularia hypogcea. 

of them will answer pretty well for them all. To those unacquainted with 
the habits of these owls, both the English and technical names might 
be misleading, giving one to suppose that the birds actually burrowed ; 
such, we believe, is never the case, they all having the like habit of re- 
sorting to the villages of many of the species of marmot squirrels, and 
occupying their deserted burrows for the purpose of nidification ; sev- 
eral families of these owls often being found in the same village. S. 
cunicularia hypogcea occurs upon the open and treeless prairies T , .est of 
the Mississippi River, where the writer has had abundant opportunity 
to study its very unique and interesting habits as well as to secure un- 
limited material for the purposes of dissection. 

We are well aware, however, that other naturalists have believed, to 
a greater or less extent, that Speotyto really, under certain circum 
stances, burrows for itself, but owing to the fact that it has never ac- 
tually been known to construct an entire burrow, or what I must believe 
any part of one, and in face of the apparent inadaptability of its small 
feet and not overpowerful beak to accomplish such a task, we can hardly 
38 H 593 







594 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



accredit the statement. This idea has no doubt been brought about by 
the fact that sometimes a few families of prairie dogs (Cynomys), for in- 
stance, will start an independent village, which, from some cause or the 
other, they are afterwards led to desert ; whereupon the owls are only 
too glad to avail themselves of the empty burrows, and the traveller is 
very likely to find one or two such peaceful colonies in the course of his 
rambles, as the writer has, where the owls are present, but all signs of 
the rodents obliterated, in some cases even the grass and flowers having 
grown again to the very entrances of the burrows. 

Speotyto, unlike the majority of the members of the great family to 
which it belongs, is not strictly nocturnal in its habits, but on the con- 
trary one may find them wide awake and active, in the villages where 
they are found, at almost any hour of the day ; indeed, as a rule they 
are quite wary, and one may often resort to ali of the stratagems his 
experience has taught him before he succeeds in securing a specimen. 
In powers of flight they are weaker than most owls, a fact largely due 
no doubt to the lack of exercise of this privilege. 

As we pass to the study of the skeleton ot this interesting species, we 
shall, no doubt, find several instances wherein it has been modified and 
received certain impressions due to the mode of life and habits of the 
owner, that we have so briefly called the reader's attention to. TVe add 
here also a cut showing some of the external characters of this bird. 




In enumerating and describing the separate bones of our subject, the 
smaller ones of the ear have not been taken into consideration, as they 
more properly come to be treated in the study of the organ of hearing - 7 
certain very small sesamoids may also, with propriety, be overlooked. 

The skull. — As a general rule, it is only in the young of the Class Ares 
that the many bones of the skull can be separated from one another; 
the majority of the primitive segments of ossification of the four ver- 
tebrae 1 that go to form this, the superior expansion of the vertebral col- 

1 Mr reader will no doubt remember that this monograph, accompanied by another 
of about equal size, upon the Osteology of EremopMla alpestris, appeared in the Bul- 
letin of the Tinted States Geological and Geographical Survey of the Territories, vol. 
vi, No. 1. Washington, February 11, 1881, being followed in the same year, September 
19, in No. 2 of the same volume, by my monographs upon the Osteology of the North 
American Tetraonidce and Lanius ludovicianus excubitorides. In one and all of these pa- 
pers I considered the cranium as composed of four vertebrae, as many of the old school 
comparative anatomists had done before me, and in adopting this theory I likewise 
adopted the nomenclature of the elements as given by Professor Owen, in his Anatomy 
and Phvsiology of Vertebrates. When I was first privileged to enter upon my ana- 
tomical studies, comparative and otherwise, this theory was the then prevailing one, 
and mv mind became imbued with its fascinating precepts, its plausibility and appa- 



shuieldt] OSTEOLOGY OF THE SPEOTYTO. 595 

umu, being firmly anchylosed together, with their sutures completely 
obliterated when the bird has attained maturity. This is eminently 
the case in the adult skull of the species we have before us, so much so, 
in fact, that, with the exception of certain bones that remain perma- 
nently free during life, we will uudertake to describe the skull only as 
it presents itself to us in the adult as a whole. In referring to certain 
points for examination, then, in this part of the skeleton, we will have 
to rely largely upon the reader's familiarity with general anatomy. ; 
and the extent and position of the bones as they occur in the variously 
shaped heads of immature birds. The major part of the occipital 
lies in the horizontal plane, only that portion which originally con- 
stituted the superoccipital segment and the posterior third of the ex- 
occipital segments curving rather abruptly upwards to meet the mas- 
toids and parietals. All its primary parts are thoroughly coalesced, 
and its articulations with the surrounding bones obliterated, save a fine 
ridge, running transversely, just anterior to the condyle, separated from 
it by a depression which seems to indicate the remains of the occipito- 
basi- sphenoidal suture. Posterior to the foramen magnum the bone rises 
and displays a well-marked "cerebellar prominence," with a depression 
on either side of it. On the summit ot this prominence, in the median 
line, just before we arrive at the foramen magnum, we find the super- 
occipital foramen. This foramen varies in size and shape in different 
individuals — in size, from one to two millimetres; in shape, from a circle 
to a transverse ellipse, though it is usually small and circular. It is 
said to be formed by a thinning of the bone due to muscular pressure 
from without and the pressure of the cerebellum from within ; in the 
fresh specimen it is covered by a thin membrane. Lying in the horizon- 
tal plane, anterior to the cerebellar prominence, is the foramen magnum. 
In shape it resembles a square with the four angles rounded off. Its 
average measurement is five millimetres transversely and four millime- 
tres antero-posteriorly, the latter diameter being encroached upon by 
the occipital condyle in the median line. The occipital condyle is sessile, 
though raised above the level of the basis crauii, hemispheroidal in form, 
with a minute notch marking it posteriorly in the middle. Immediately 
beyond the condyle appears a depression, on either side of which are 
seen the precondyloid foramina for the transmission of the hypoglossal 
nerves; they are extremely small, and open anteriorly. External to 
these, lying in the same line transversely, is seen a group of usually 
three foramina for the passage of the glossopharyngeal and vagus 

rent universal adaptability; in fact, its truth became one of the treasured results of 
my university education. Several years elapsed before I was again allowed to renew 
my favorite study, during wbich time the vicissitudes of my life allowed but little op- 
portunity to follow the many advances in this important science, and when the day 
finally came, and the above monographs were written, and I drew the plates illustrating 
them, I was many hundreds of miles, and for several years, removed from all that one 
has access to in large cities and scientific circles. From this standpoint they must be 
judged, then, and for this reason did errors and theories creep into them that are " so 
dangerous to the credit of comparative anatomy." It is believed that the Osteology of the 
Catliartidce will be largely exempt from sucb errors. The author deems it entirely un- 
necessary to enter upon the merits or demerits of any theory here, simply announcing 
in connection with what has already been said above, that he believes the vertebral 
theory of the cranium to be untenable in the light of modern science, and incompat- 
ible with modern thought. Aside from the theory involved, it is hoped t hat the purely 
anatomical facts will be found to be correct as given, and as any attempt to eradicate 
the theory from these memoirs would simply result in a rewriting of the whole, the 
author has allowed them to stand substantially as they first appeared, simply offering 
his reader the. above explanation and making such changes as he deems best through 
the medium of foot-notes; introducing into the body of the paper only such material 
and facts as he has been able to gather since, bringing the whole up to the standard of 
a revised edition. 



59G GEOLOGICAL SURVEY OF THE TERRITORIES. 

nerves and the internal jugular vein. The lateral terminations of 
•the occipital, the paroccipital processes, are large, thin, pointed for- 
wards, and on a lower level than the rest of the bone, forming a 
large part of the floor of the tympanic cavity. The semi-elliptical 
contour of the cranium, regarding it from a basal view, is well car- 
ried out laterally by the wing-light and attenuated mastoids. They 
contribute largely to the formation of the walls of the tympanic 
cavity internally, and externally assist in some degree towards com- 
pleting the temporal fossae. These fossae are deep ; commencing 
posteriorly on either side at the external borders of the depressions 
already mentioned that bound the cerebellar prominence laterally, they 
take a course upwards and outwards, terminating at a foramen that lies 
just within the posterior periphery or the orbit, which foramen allows 
the passage of the tendon of the temporal muscle. From the upper 
boundary of the temporal fossae to where the frontals suddenly abut 
against and even overhang, to some extent, the nasals, the external and 
superior surface of the skull is of a pearly whiteness and very smooth in 
the dry skeleton, presenting not a trace of the sutures between the bones 
that go to form it, the frontals and parietals. This surface is divided 
by a well-marked furrow, that extends in the median line between the 
cerebellar prominence and the upper mandible. It is deepest in the 
parietal region. Close inspection of this area reveals minute ramifying 
grooves for the lodgment of vessels, one set running in the direction of 
the temporal fossa? and another toward the orbits. In the " bird of the 
year" the skull cap is very thin and brittle in the dry condition ; but a 
very different state of affairs presents itself when we remove a section 
of the cranial vault from above, in the adult, where the skeleton is full- 
grown, such as we have before us. We find exposed to our view one of 
the common characteristics of the family ; the two tables are light, thin, 
but compact, with a goodly supply of diploic tissue between them, at- 
taining a thickuess in some localities, notably above the exit of the olfac- 
tories, of two millimetres or more. Owing to the large orbital cavities, 
the brain-case is crowded to the rear to such an extent that the fossae 
for the cerebral hemispheres are situated immediately over the cavities 
intended for the other encephalic lobes. We find the internal opening 
of the foramina, already described, at the base of the brain. The petro- 
sals have the appearance of two white leaves, harder than the surround- 
ing bone, slightly turned upon themselves, with their stems leading 
towards the fossa for the hypophysis. They present for examination the 
openings for the portio dura and portio mollis, the former foramen being 
ou a lower level and anterior to the latter. In the median line running 
from the cerebellar fossa to the exit of the first pair of nerves along the 
roof is a raised crest, grooved on its summit for the longitudinal sinus. 
It sinks for a little distance, in the fresh specimen, into the cerebral in- 
terspace. The " sella turcica" is deep, its long axis being perpendicular 
to a plane passing through the foramen magnum. It has at its base the 
openings for the carotids. Immediately beyond its anterior superior bor- 
der is seen the optic groove, with its foramen at either end, for the passage 
of the optic nerves and lodgment for the optic chiasma. Above the optic 
foramina, situated still more anteriorly, is a conical pocket, pointing for- 
wards and a little upwards, with the olfactory foramina at its apex, two in 
number, giving passage for the nerves to the orbits. The bast sphenoid is 
thoroughly united with all the bones it comes in contact with, except the 
pterygoids, palatines, and tympanies. Its anterior process — the basi- 
presphenoid — lost-s itself in the interorbital septem, not a trace re- 
maining of the original margins of the two bones. Its wings, the orbito- 



shufeldt.] OSTEOLOGY OF THE SPEOTYTO. 597 

sphenoids and the ali-sphenoids share the same fate with the bonesthat 
surround them. They form the larger part of the posterior wall of the 
orbital cavities. With the bod}* of the bone the ali-sphenoids assist in 
closing in the tympanic cavities. We cannot positively state that this 
owl possesses a true boney vomer. The -- pterapophysial" processes of the 
basi-sphenoid are present; they are short, thick, and elliptical on section, 
crowned by facets of the same figure at their distal extremities, which 
look downwards, forwards, and outwards, articulating with a similar 
facet at the middle third and posterior border of each pterygoid. The 
bone also presents for examination the usual nervous and arterial foram- 
ina and grooves for the Eustachian tubes, the foramina being particu- 
larly worthy o£ notice on account of their marked individuality, all of 
them being distinct and nearly circular. The tympanies are free bones, 
and carry out all the usual functions assigned to them. The mastoid, 
condyle is long, affording by its extension an additional margin at the 
under side at the end of the bone for attachment of the ear-drum; the 
neck between it and the orbital process is somewhat constricted, and 
presents a large pneumatic foramen on the inner surface. The pointed 
orbital processes extend upwards, forwards, and in wards, slightly clubbed 
at their extremities ; they project into the space half way between the 
pterygoid and wing-like post-frontal. The mandibular condyle is double ; 
the inner one is a semi-ellipsoid, placed transversely ; the outer an irreg- 
ular figure, and separated from the inner by a shallow pit. The oval, 
cup-shaped cavity for the reception of the tympanic extremity of the 
squamosal looks directly forward. Between the orbital process and inner 
mandibular condyle, on the free edge of the bone, is seen a small artic- 
ular surface for the tympanic extremity of the pterygoid. The pterygoids 
diverge from each other towards the tympanies by a very open obtuse 
angle. They are slender and scale-like, being compressed from above 
downwards, twisted on themselves at their tympanic extremities, caus- 
ing the long axis of the articular facets for the articulation with these 
bones to be vertical. As already described, they have a mid-posterior 
facet, which meets the pterapophysial process of the basi-sphenoid. An- 
teriorly they do not touch each other, but articulate with the extremities 
of the palatines, and the combined four bones touch, and in the living 
bird glide over for a limited distance the lower border of the rostrum of 
the basi-presphenoid. The anterior ends of the palatines articulate by 
an anchylosed schindylesial articulation between the lower surfaces of 
the inaxillaries and the thin upper surface of a bony process extending 
backwards from the intermaxillary. From this point they slightly di- 
verge from each other and become broader, being broadest about their 
middles ; they then rather abruptly approach each other posteriorly, where 
they form the joint with the pterygoids already described. Their pos- 
terior ends are kept slightly apart by the lower border of the presphe- 
noid. They are flattened from above downwards throughout their en- 
tire extent. Their outer borders are sharp, and form from one end to 
the other a long convexity. As the inner and concave borders ap- 
proach each other posteriorly they develop a raised rim on their under 
sides, thereby affording a greater surface for muscular attachment. 
Above, near their middles, they aid the maxillaries (and in large part de- 
veloped from them) in supporting on either side an irregular spongy bone, 
that serves the double purpose of narrowing the apertures of the poste- 
rior nares and adding bony surface to the roof of the mouth by constric- 
tion of the palatine fissure. 1 As is the rule in nearly all birds, the tym- 

1 These are the maxillo-palatines of Huxley, and the reader is referred to that au- 
thor's invaluable paper upon the Classification of Birds, etc. Proc. Zool. Soc. Lond , 
1867, p. 441, where these bones are shown in Otus vulgaris, fig. 26. 



598 GEOLOGICAL SURVEY OF THE TERRITORIES. 

panic end of the infraorbital bar is on a lower level than the maxillary 
extremity 5 it is received into the cup-like articulating cavity on that 
bone. The two oblique sutures, persistent in many birds, and denoting 
the original division of this bony style into three separate bones, the 
maxillary, malar, and squamosal, are here entirely effaced. As a whole, 
it is compressed from side to side, and of ample size in comparison with 
other bones of the head. At about the locality of the malo-zymotic 
suture the bone throws upwards a thin expansion that meets the de- 
scending postfrontal, thus completing the orbital circumference at that 
point. Its anterior and fixed extremity is made up by the maxillary. 
Here it forms externally a portion of the posterior surface of the bill, 
while internally it assists in forming the roof of the mouth and floor ot 
the nasal cavities, and otherwise behaves as already described. The 
lacrymals are extremely spongy in texture, covered by an outside delicate, 
compact bony casing. They articulate above by a ginglymoid joint with 
the posterior border of the nasals, resting below on the spongy bones de- 
veloped from the superior surfaces of the maxillaries. They are limited 
to a slight movement inwards and outwards, and aid in separating the or- 
bital cavities from the rhinal chamber. Externally they present for ex- 
amination a shallow groove traversing the bone obliquely downwards and 
forwards and a little inwards for the lacrymal duct. The orbital cavities 
are very large, and remarkable for the completeness of their bony walls 
and the near approach their peripheries make to the circle, any diam- 
eter of which measures the merest trifle above or below two centimetres. 
The septum in the adult bird has rarely more than one small deficiency 
of bone in it. This usually occurs in about the position shown in PI. I. 
The sutures among the various bones have entirely disappeared, nothing 
being left to define the exact outline of the vomer especially. The groove 
for the passage of the olfactory nerves forward is well marked, the cra- 
nial foramina for them being distinct, one in each orbital cavity. This 
also applies to the openings for the optic nerves. The extent of the roof 
is increased on either side by a superorbital process (shown in PI. II, 
Fig. 1) that points dowmvards, backwards, and outwards, and serves for 
membranous attachment. The posterior walls are marked by ramifying 
grooves for vessels. They have a direct forward aspect, which is en- 
hanced b3 T the low descent of the broad and thin postfrontals. Ante- 
riorly, the aperture between these and the rhinal vacuities is diminished 
by the lacrymals externally and by a wing-like plate thrown off from 
the prefrontal internally. This latter bone here terminates in a sharp 
concave border, with a descending ridge on either side just within it. 
The floors of the orbits are more complete than is usually seen in the class, 
due to the flatness and position held by the pterygoids and palatines, 
the wing like process of the ethmoid just referred to, and the pterapo- 
physial processes of the basi sphenoid. The sclerotals number from 
fifteen to sixteen, all of them being about the same length, but varying 
as to their width ; in figure they are trapezoidal and universally oblong, 
with the short parallel side in the circumference of the cornea and the 
opposite one resting in the periphery of the posterior hemisphere. We 
have never observed one that was wide enough to appear square. They 
are rather thin, concave outwards, very slightly movable at their op- 
posed edges, and carry out their usual function of maintaining the form 
of the optical apparatus. The upper mandible of this bird is made the 
more conspicuous and distinct from the remainder of the skull by the 
abrupt way in which it is attached and the much firmer texture of the 
bone. The mandibular culmen is perfectly convex from the tip of the 
sharp-pointed extremity to where it suddenly terminates under the 



shufeldt.] OSTEOLOGY OF THE SPEOTYTO. 599 

slightly overhanging* frontals, or, more correctly, the minute surface ap- 
pearance of the prefrontal, for although it is not evident in the adult that 
that bone makes itself visible at this point, yet it may be demonstrated 
in skulls of younger specimens. The culmen, as in other birds, is formed 
by the intermaxillary, which is here firmly united with the nasals, and 
the two in conjunction form the peripheries of the truly elliptical external 
nasal apertures or nostrils, the first bone bounding them anteriorly, 
while the latter completes their arcs in the rear. These in the dry skull 
measure through their major axes seven millimetres, and through their 
minor ones barely five millimetres. They have a distinct ring raised 
around their circumference, which is wanting, however, where they 
nearest approach each other anteriorly at the culmen. The plane of the 
nostril faces upwards, outwards, and forwards ; the nostrils are com- 
pletely separated from one another by a vertical bony septum, developed 
from the intermaxillary, not a common occurrence in birds. They have, 
in addition, a concave bony floor, that rises behind into a posterior wall 
leaving really two semicircular openings just beneath the culmen, sepa- 
rated from each other by the vertical septum. The osseous mandibular 
tomium, also a part of the intei maxillary, is as sharp as when the bill is 
sheathed in its horny integument. The arc is concave, and falls off 
rapidly as it approaches the tip of the beak. Occasionally, in very old 
birds, the eUimo-turbinal bones in the nasal passages may ossify. The 
nasals form here the sides of the bill, and are firmly anchylosed to the 
bones they meet, except the lacrymals. The movability of the frouto- 
mandibular articulation is limited. The dry skull is extremely light and 
brittle, giving one the sensation in handling it that he might experience 
while examining an egg from which the contents had been removed. A 
line drawn from the tip of the upper mandible to the outermost point of 
one tympanic, around the arc of the cranium to a similar point on the 
opposite side, and back to the point of departure, describes nearly the 
sector of a circle. The longest radius, which is in the median line, meas- 
ures four and one-half centimetres, the cord between the tympanies 
about three centimetres. 

The hyoid arch. — The hyoid arch is suspended from the base of the 
skull by its usual attachments. In this Owl it consists of but six very 
delicate little bones, involving five articulations. The tips of the up- 
turned posterior extremities are about opposite the lower borders of the 
temporal fossae, its two limbs diverging from each other at an angle equal 
to that made by the lower mandible. The cerato-hyals are rathe r large 
in comparison with the other bones. They are joined both anteriorly 
and posteriorly by bony bridges, forming a fenestra between them, to 
be filled in by a thin membrane. The amount of divergence they make 
from each other is less than that made by the hypo-branchial elements 
of the thyro-hyals. Anteriorly, the bone connecting them supports a 
cartilaginous glosso-hyal, while the posterior connection presents for 
examination the usual smooth articulating surface that enters into the 
arthrodial joint it makes with the basi-hyal. The basi-hyal and uro- 
hyal are confluent, not a sign of the point of union remaining. The 
latter bone is continued a short distance posteriorly by a tip of cartilage. 
The anterior end of the basi-hyal is devoted to the articular surface for 
the bone connecting the cerato hyals, forming the joint mentioned above. 
It is concave from above downwards, convex from side to side, the lower 
lip being the longer. It will be plainly seen that this combination grants 
to the tongue a movement in the vertical and horizontal planes. The 
anterior articulating heads of the hypo-branchial elements of the thyro- 
hyals are opposite each other, each being received into the diminutive 



600 GEOLOGICAL SURVEY OF THE TERRITORIES. 

acetabulum intended for it at the side of the united basi- and uro-hyals, 
and most probably at the junction of the two latter bones. These two 
elements are long bones having a cylindrical shaft, terminating at either 
end in an articulating head. They are the longest bones in the hyoid arch, 
andhaveagentlecurvatureupwardstliroughout their extent. Their inner 
heads form an arthrodial joint on either side with the outer heads of 
the cerato-branchial elements of the thyro-hyals. These, the last bones 
of the arch, are joined in the manner already shown above. Their inner 
ends are quite pointed, even as far as the bone goes, the extreme points 
being finished off with cartilage. They curve upwards from about their 
middle thirds, and, like the first elements of the thyro-hyals, they are 
long bones, but with curved cylindrical shafts, the outer end, however, 
being the only true articulating one. 

The lower mandible — (Pis. I and II, Fig. 3). — That portion of the bone 
which originally was separate as the dentary element, and as far back as 
to include the interangular vacuity, is firm and compact, while the re- 
mainder has much the same character as the bones of the cranium, being 
cellular and light, having only a very thin outside layer of the harder 
tissue. All of the primary segments ar > firmly knitted together, the only 
sutural trace to mark the margins of any 6ue of them being the posterior 
border of the dentary elements as they bound the fenestra before and 
slope away beneath it. The articular extremities are some little distance 
below the upper outline of the bone. Their superior surfaces are in- 
dented so as to accurately receive the condyles of the tympanies on 
either side, forming the joint that allows the opening and closing of the 
mandibles. Their under surfaces are smooth and rounded, having a 
fine ridge running across them transversely. Internally they are drawn 
out gradually into subcylindrical processes that point upwards, in- 
wards, and a little forwards, exhibiting superiorly on each, about the 
middle, an oval pneumatic foramen. The upper edge rises rather ab- 
ruptly from the articular ends, presenting as it arrives near the general 
level a rudimentary coronoid for the insertion of the tendon of the tem- 
poral. With the exception of a little elevation where the dentary ele- 
ment meets the surangular, the superior outline is unbroken; it falls 
away rapidly as it approaches the symphysis, where, with the opposite 
border, it completes a little notch at the extremity. The tomium is 
not as sharp as in the upper bill, and the mandibles do not fit nicely to 
each other until covered with their horny sheaths. The inferior border 
is rounded throughout its extent, and on a level at its posterior com- 
mencement with the under surfaces of the articular ends and running 
nearly parallel with the superior. The curve described by the rami 
before they meet at the symphysis interiorly approaches the parabola in 
outline. The sides of the jaw are nearly smooth internally aud exter- 
nally. The vacuity that occurs in so many birds at the junction of the 
middle and inner thirds is rather large, long, and spindle-shaped, and 
filled in, in the fresh state, by an attenuated membrane. 

Professor Huxley, in his Classification of Birds (Proc. Zool. Soc. 
Lond., 1867, p. 462, 5 JEtomorphw), presents us with some of the most 
important cranial characters of the Strigldce; and we find through the 
literature of the subject not a few authors who have touched upon the 
osteology of this interesting group of birds. The attention of ornitholo- 
gists and others has been directed on several occasions to the asym- 
metry occurring in the skull of certain species of Owls, notably in Nye- 
tale. In 1870, Dr. T. H. Streets, of the United States Navy, noticed 
this point and published his observations. (Proc. Acad. Nat. Sci. Phil , 
1870, p. 28.) In the next year, Kobert Collett, esq., of Norway, noted 



bhufbldt.] OSTEOLOGY OF THE SPEOTYTO. 601 

the same thing in N. tengmalmi. (P. Z. S., 1871, pp. 739-743.) Figures 
showing this condition are given by Mr. Collett in an interesting paper 
of his that he kindly sent me upon the Craniets og Oreaabningernes 
Bygning hos de nordeuropaeiske Arcer af Familien Strigidae. Mr. Ridg- 
way has likewise figured the skull of Kyctale richardsoni in the North 
American Birds. In TJlula cinerea we have another variety of cra- 
nial asymmetry; in the specimen before me the post-frontal wing is 
thrown farther outward on the right side; this is not the case in Strix 
nebulosa, a species that has a perfectly symmetrical skull. This charac- 
teristic occurs in other Owls. In Surnia funerea we find the osseous 
nasal septum well perforated, at its upper and inner part, very much as 
it is in Circus. The Hawk Owl has likewise superorbital processes of 
the same form as those we have described above for Speotyto. The 
periphery of the orbit above in Surnia is slightly rounded, not nearly 
as much though as it is in Strix nebulosa, much less than in Asio. These 
borders are very sharp in Scojjs, while in Aluco they nearly merge into 
the orbit. This latter Owl has a skull that is at once strikingly differ- 
ent from other forms of the family, being long and. narrow, the orbits 
being separated, by a great, thick, spongy septum, the wings of the 
ethmoid are likewise spongy cylindrical masses, and. the lacrymals are 
very large, being composed of the same material. For a family where 
the skulls of the various forms vary so much, differ so much from each 
other, we note quite an exception in the crania of Nyctea and Bubo, 
owls that have skulls strikingly alike, except in point of size, and a few 
minor differences. 

The spinal column; cervical portion. — There are fourteen cervical ver- 
tebras, each one having a more or less free movement with the one be- 
yond and behind it, maintaining in all positions some variation of the 
usual sigmoid curve observable in the division of the vertebral column 
throughout the class. The arrangement, as well as the direction, of the 
planes of the zygapophysial articular surfaces allow considerable rotary 
movement and bending in the vertical plane, with combinations of the 
two. It is a common habit of this bird, among other of his antics, to 
duck his head smartly downwards and again upwards, several times in 
succession, upon being approached. The relative position of the cervi- 
cals has been figured in Plate I from the dead bird, placed in the act of 
this particular manoeuvre, in a specimen after careful dissection. The 
calibre, as well as shape, of the neural canal in this portion of the spinal 
column varies at different points. It originates at the atlas as a trans- 
verse ellipse, with a major axis of four millimetres and a minor axis of 
a little less than three millimetres ; this is about the maximum capacity 
throughout the entire canal. From the atlas to the sixth or seventh 
vertebra the ellipse gradually approaches the circle, with a marked 
diminution in size, its diameters being at the seventh about two milli- 
metres in any direction. From this point to the twelfth, inclusive, it 
rises as it fell from the atlas, and in the same manner, when we again 
discover a transverse ellipse, perhaps a jot smaller than the one de- 
scribed in speaking of the atlas. In the thirteenth the canal is smaller 
than, though in all other respects resembles, the twelfth, but an abrupt 
change takes place in shape as we pass to the fourteenth or last cervi- 
cal, where the form of the neural tube suddenly approximates the cir- 
cularity of the dorsal vertebra?. The vertebral canal begins, circular, on 
either side at the third cervical vertebra, most of its length being im- 
mediately beneath the prezygapophyses of each segment. It is formed 
in the usual manner by the di- and par-apophysial processes uniting 
laterally with the pleurapophysial elements. Sinali at the cephalic 



G02 GEOLOGICAL SURVEY OF THE TERRITORIES. 

extremity of the column, its calibre gradually increases in each ver- 
tebra as we proceed toward the thoracic extremity, until it attains its 
maximum capacity at the eleventh vertebra. In the twelfth the in- 
tegrity of its walls is lost by a parting of the par- and pleur-apo- 
physial elements, with a disappearance of the former, leaving it no 
floor, so that in this vertebra it ceases to be a closed canal. The most 
prominent object presenting itself for examination in the atlas, superi- 
orly, is the deep reniform cavity for articulation with the occipital con- 
dyle of the basi-cranii. It makes up to the entire superior articulating 
surface of what would first appear to be the centrum of this vertebra, 
unless we should not consider such to be the case until the odontoid 
process of the vertebra next below, the true centrum of the atlas, lends 
its assistance, in which event the surface of this articulation is only 
complete when made so by the extremity of the process just alluded 
to. A membrane sometimes stretches across this interspace, sepa- 
rating the extremity of the odontoid from the condyle ot the occi- 
put; this is not invariably the case, however, as in many of the 
individuals we have examined a minute vacuity exists, allowing the 
process to come iu immediate contact with the condyle at one point. 
Below and posteriorly there is another articulating surface, convex for 
the centrum of the axis and concave for its odontoid process, accurately 
meeting the opposed surface of this vertebra and forming the atlo-axoid 
articulation. A lip of bone, a portion of the hypapophysis of the verte- 
bra we are now describing, projects downwards and shields this joint in 
front, overlapping, indeed, a good part of the axis. The neurapophyses 
of the atlas are slight in structure. The concave postzygapophyses ar- 
ticulate with the convex prezygapophyses of the axis. The bone is de- 
void of a neural spine. In the axis we find both an hypapophysis and 
neural spine developed, the former being produced from the ridge on the 
anterior aspect of the centrum of the bone. The odontoid process arises 
vertically from the posterior margin of the upper surface of the centrum. 
Its summit and anterior face are convex and articulating, while behind 
it is flat and continuous with the spinal canal. The facet for articula- 
tion with the centrum of the third vertebra looks downwards and in- 
wards, is convex from side to side and concave in the opposite direction. 
The postzygapophyses are concave, look downwards and outwards, the 
conditions in the prezygapophyses being exactly the opposite ; this is 
the rule throughout the cervical portion of the columu. After we pass 
the atlas and axis, we find in the third cervical vertebra, here, as in 
most vertebrates, parts that are common to the series of this portion of 
the column, deviating but slightly from each other as we examine them 
in seriatim; but gradually as this deviation proceeds, some requisite con- 
dition is brought about when the climax is attained. The fact of the 
presence of a neural spine on the axis is conveyed, though in a less 
marked degree, to the third or next vertebra below, where it occupies a 
position about in the middle of the bone. As we descend, this process 
becomes less and less prominent, being found set further back on each 
successive vertebra; it disappears about entirely at the tenth, after 
which it rapidly begins to make its appearance again, assuming its former 
position in the middle of the vertebra, being quite evident in the twelfth 
in the shape of a pointed spine, while in the fourteenth it bears the quad- 
rate form, with extended crest, being the first step towards an assumption 
of that notorious feature found further on in the dorsals. In the third 
vertebra the space betweeu the pre- and postzygapophyses is almost en- 
tirely filled in, a minute foramen on either side alone remaining, by a 
lamina of bone extending from one process to the other, giving to this 



shufeldt.1 OSTEOLOGY OF THE SPEOTYTO. 603 

vertebra a much more solid appearance, which in reality it possesses above 
that attained by any of its fellows. This bony lamina is reduced in the 
fourth vertebra to a mere "interzygapophysial bar" connecting the pro- 
cesses, while in the next succeeding one or two vertebrae it occurs only 
on the prezygapophyses more as a tubercle, being directed backwards, 
then disappearing entirely, is to be found again only on a few of the 
last cervicals as an ill defined knob, still retaining its original position- 
The diapophyses at first project nearly at right angles from their re. 
spective centra, then approach the median line by being directed more 
backward near the centre of the cervical division of the column, and on 
Hearing the dorsals again gradually protrude more and more directly 
outward. The prezygapophyses of the ninth cervical support well- 
marked anapophysial tubercles, which are feebly developed also on a 
vertebra or two both above and below the ninth. The joints between 
the bodies of the cervicals of this Owl are upon the same plan as those 
found throughout the class; the anterior facet being concave from 
side to side, convex from above downwards, the reverse being the case 
with the posterior facets, and when articulated fitting accurately into 
each other. The pleurapophysial elements, well marked in all the cervi- 
cals after passing the axis, become in the thirteenth vertebra a free 
cervical rib, about three millimetres in length, without neck or true 
head, being merely suspended on either side from the diapophysis of 
the vertebra, and freely movable on its exceedingly minute articulating 
facet. 

Attached to the last cervical we find the second pair of free pleura 
pophyses, about two thirds as long as the first pair of dorsals or true 
ribs of the thorax, terminating in pointed extremities and articulating 
with the vertebra by both capitula and tubercula, the former on ellipti- 
cal facets, placed vertically on either side of the centrum at the anterior 
margin of the neural canal, and the latter on rounded facets beneath 
the diapophyses. The tubercle on one of these ribs is nearly as long as 
the neck ; at the junction on the posterior side is found a pneumatic 
foramen of considerable size. These ribs are more or less flattened 
above from before backwards, being convex anteriorly, concave poste- 
riorly, becoming rounded below. From the third to the ninth vertebra, 
inclusive, appear beneath the vertical canal anteriorly well-developed 
styliform parapophysial processes, directed backwards and downwards. 
They are best marked on the segments of the middle of the neck. There 
is no instance in this bird of these processes being produced so far 
backwards as to touch the next vertebra below ; their tips, as a rule, 
about overhanging the middle of the centrum of the vertebra to which 
they belong. We have found in specimens of Bubo virginianus the 
parapophyses of the fourth vertebra overlapping and touching the fifth 
for a millimetre or more. The third and fourth cervicals have, beneath 
in the median line posteriorly, strongly developed hypapophyses, quad- 
rate in form, a process that exhibits itself on the fifth vertebra, ante- 
riorly, merely as a small tubercle. On the sixth this tubercle has disap- 
peared, and has been supplanted by two others that are now found just 
within the periphery of the anterior facet of the centrum on the para- 
pophysis of each side, beueath and inclined toward each other. These 
processes, now a double hypapophy sis apparently developed from the par- 
apophyses, continue to increase in size and inclination towards each other 
on the next three vertebras, so that on the ninth, where they last appear, 
they nearly form a closed canal. The passage between them is intended 
for the carotids, to which they afford protection. The hypapophysis of 
the tenth, eleventh, and twelfth vertebras is single, large, quadrate, and 



604 GEOLOGICAL SURVEY OF THE TERRITORIES. 

directed forwards and downwards. There are three on each of the last 
two vertebrae, each having an independent root, the two lateral ones 
directed downwards, forwards, and outwards, with characteristics similar 
to the one in the median line. Several pneumatic and nutrient foramina 
X>erforate each cervical vertebrae at various points, except in the axis 
and atlas, where, after diligent search, aided by the lens, we have sig- 
nally failed to discover them. 

Dorsal vertebrce ; vertebral and sternal ribs ; sternum. — The dorsal ver- 
tebrae number five 5 the anterior one articulates with the last cervical 
and the last dorsal with the first sacral. Although the dorsals of this 
biid fit very snugly to each other, it requires no further maceration to 
separate them from one another than it does to remove the ribs from 
their attachments. This close interlocking, however, greatly diminishes 
the movement of this division of the spinal column, bestowing upon it a 
rigidity only exceeded by the anchylosed vertebrae of the sacrum ; yet, 
it must be understood, they do eujoy, in this Owl, a considerable degree 
of movement, especially laterally. The neural spines have here attained 
their maximum development, forming, when taken together, an elevated 
and compressed median crest, with a thickened summit, and having a 
firm hold upon the remainder of the vertebrae below. Taken separately, 
the last is the smallest, the fourth next, the first next, and the second 
and third the largest. Their anterior and posterior borders are concave, 
allowing, when articulated, spindle-shaped apertures to exist among 
them, while their summits are produced backwards and forwards, thick- 
ened, and wedged into each other. This wedging is performed in the 
following manner: Tbe posterior extremity of the crest forming the sum- 
mit of the neural spine of the first dorsal divides and receives the an- 
terior extremity of the crest of the second. This same arrangement ex- 
ists between the second and third, and at the summit between tbe third 
and fourth, but the fourth immediately below the junction also divides 
for a little distance and receives the edge of the posterior rim of the 
third, just beneath the union of the crests. This latter method of join- 
ing is feebly attempted between the fourth and last. (See PI. I.) The 
neural canal is nearly cylindrical in the dorsal region, its calibre being 
less at the sacral extremity, and rather compressed from side to side, as 
are the centra as we approach that end, each one being a little more so 
than its neighbor beyond. Viewing these five vertebrae from above in 
the articulated skeleton, we observe the spinous crest already described ; 
Ave are struck with the regularity with which the postzygapophyses over- 
lap and adjust themselves to the prezygapophyses from before back- 
wards, like the scales in some fishes, the facets of the former facing 
downwards and outwards, the opposed surfaces of the latter facing up- 
wards and inwards. The neurapophyses are horizontally compressed 
and rather broad ; the diapophyses jut from them at right angles from 
points about their middles. There is an inclination for the latter to 
be directed slightly backwards as we near the sacrum. The diapophy- 
sis of the first dorsal is the shortest and stoutest, that of the last the most 
delicately constructed. Superiorly, these processes support metapophy- 
sial ridges at their extreme outer borders. These ridges on the diapophy- 
ses of the first dorsal are the largest, rounded at both ends, extending 
a little both backwards and forwards, but far from touching the ridge 
either in front or behind them. The metapophysials of the last dorsal 
are smaller, sharp, styliform, and project only forwards, though they 
do not by any means touch the diapophyses in front of them. On 
the intermediate vertebra they change gradually between these two 
extremes, but in no instance meet the diapophyses of the vertebra be- 



bhufeldt.] OSTEOLOGY OF THE SPEOTYTO. 605 

fore or behind them, and thus constitute an additional aid to the rigid- 
ity of the back, as it does in other species of this family and in many 
other birds. The centra increase in depth beneath the neural canal the 
nearer they are to the sacrum. In the first dorsal the body measures 
about one millimetre, the vertical diameter of the canal being three; in 
the last dorsal it equals the diameter of the canal. The interarticular 
facets are in the vertical plane, with their concavities and convexities op- 
posed to each other, as they were described when speaking of the last cer- 
vical vertebrae. The bodies are about of a length, constricted at their mid- 
dles and expanding towards their extremities. The first two dorsals each 
bear in the median line, beneath, an hypapophysial process of consider- 
able size, affording abundant surface for attachment of some of the muscles 
of the neck. The process of the first dorsal has one common trunk, with a 
compressed midprong ana two lateral and pointed subprocesses. (See 
PL II, Fig. 5.) The second dorsal possesses a single long hypapophysis, 
quadrate in form, dipping into the chest further than the first. There 
is not a trace on the remaining dorsals of this appendix. Parapophysial 
processes, so prominent in nearly all the cervicals, afford in the dorsal 
vertebrae- simply articulating facets for the capitula of the pleurapophy- 
ses situated just within the anterior margin of the neural canal of each 
centrum, never extending to the vertebrae beyond, forming the demi-facet 
of andranatomia. Immediately above these facets, on either side, may be 
noticed a group of pneumatic foramina of various sizes and shapes, and 
again, anterior to these foramina, the rim of the body of the vertebra for 
a limited distance becomes sharply concave, being opposite to a like con- 
cavity in the next vertebra, the two, when opposed and articulated, form- 
ing the oval foramen for the exit of the dorsal nerves. Elliptical artic- 
ulating facets for the tubercula of the pleurapophyses, looking down- 
wards and outwards, are seen on the inferior ends of the diapophyses 
with a midridge running from each facet to the base of the process, to 
be expanded and lost on the sides of the centra. As there are five dor- 
sal vertebrae, so are there live pleurapophyses articulating with them ana 
with the haemapophyses below. Each rib is attached to a single verte- 
bra, as shown while speaking of the dorsals. The necks of these ribs 
become more elongated the nearer they are to the pelvic extremity of 
the body, the first possessing the shortest. This is exactly reversed in 
regard to the pedicles bearing the tubercula, being the longest in the 
first pleurapophysis and shortest in the last. This contraction of the 
pedicles is progressively compensated for by the lengthening of the cor- 
responding and respective diapophyses of the vertebra to which they 
belong. Viewing the ribs from the front, in the skeleton, the curve they 
present resembles the quadrant of a shortened ellipse, the vertex of the 
major axis being situated at the base of the neural spines; viewed lat- 
erally, the curve issigmoidal, though a much elongated and shallow one, 
with the haemapophysial extremity looking forwards and the facet of 
the tubercle backwards. The first rib is the shortest and generally, 
though not always, the broadest ; the last being the longest and most 
slender, the intermediate ones regularly increasing in length and dimin- 
ishing in breadth from the first to the last. In form, the ribs of this 
Owl are flattened from side to side, widest in the upper thirds, narrowest 
at their middles, and club-shaped at their lower extremities, where they 
articulate with the sternal ribs by shallow facets. On the inner surfaces 
we find the necks produced upon the bodies as ridges, running near their 
anterior margins and becoming lost at about the junctiou of the upper 
and middle thirds in the body of the rib. Pneumatic foramina, from 
two to three in number and of considerable size, are found just within 



606 GEOLOGICAL SURVEY OF THE TERRITORIES. 

the commissure between neck and tubercle, posteriorly. All the verte- 
bral ribs bear a movably articulated epipleural appendage, each resting 
in a shallow cavity designed for it upon the posterior borders. They 
leave the rib at right angles, but soon turn upward with a varying ab- 
ruptness. The appendage of the first rib is situated lowest of any on 
its rib, that of the last the highest ; the facets of the others are found 
in the line joining those of the first and last. They all make acute 
angles with the bodies of theribs to which each belong, above their points 
of insertion. The angle made by the last is the least, and it increases 
to the last. The epipleurals of the leading pleurapophyses are the widest 
and generally the longest (the one on the second rib in a skeleton of 
this bird now before me is as wide as the rib at the point from where it 
starts), the one on the last rib being always the smallest. 

Clubbed at their superior extremities, each one overlaps the rib behind 
it, and in this manner add stability to the thoracic parietes, which is un- 
doubtedly one of the functions these little scale-like bones were intended 
to fulfill. The hcemapophyses connect the vertebral ribs with the sternum. 
There are six of them, one articulating with each vertebral rib and hav- 
ing a concave facet to receive it, while the last meets the sacral rib 
above and articulates with the posterior border of the fifth below. The 
first one is the shortest the most slender of all ; the fifth is the long- 
est. With the exception of the last, their superior ends are enlarged 
and compressed from side to side, while below their middles they be- 
come smaller; then turning upon themselves, suddenly enlarge again, 
so as to be flattened from before backwards, when each terminates by a 
transverse articular facet for articulation with the haemal spine. Quite 
an interspace exists between their points of contact with the sternum. 
They all make a gentle curve upwards just before meeting their respect- 
ive ribs. The hsemapophysis that articulates with the sacral rib is in- 
serted in a long, shallow groove on the posterior border of the sternal 
rib that articulates with the last dorsal pleurapophysis, but does not 
meet the sternum — simply terminating in a fine point on the posterior 
border of the sternal rib mentioned. From before backwards the ster- 
nal ribs make a gradually decreasing obtuse angle with the vertebral 
ribs, while the angle they make with the sternum is a gradually increas- 
ing acute from the fifth to the first. On the anterior surfaces of their 
expanded sternal ends are to be found on each a minute pneumatic fora- 
men or two. The anterior third of the lateral borders of the sternum 
is the £,pace allotted for the insertion of these bones. 

The Burrowing Owl being a bird not possessed of any considerable 
power of flight, a circumstance arising from the life it was destined to 
lead, or the necessity of having that flight ever long sustained, we would 
naturally expect to find, in the course of a study of its anatomy, those 
characteristic modifications of the various systems which pertain to spe- 
cies of the class in which that gift has always been a secondary consid- 
eration. Nor are we disappointed in this expectation, for a single 
glance at the size of the sternum of this Owl, when compared with the 
remainder of its skeleton with regard to areas for muscular attach- 
ments, reveals to us the disproportion of the surface supplied by that 
bone for the attachment of the pectorals. That its dimensions are rel- 
atively contracted is proved by actual, comparative, and proportional 
measurements of the bones with other species of its family, individuals 
of which, at the best, are not noted for their powers of flight, and con- 
sequently the sternum does not present so prominent a feature of 
the skeleton as it does in other species of the Class Aves where vigor- 
ous flight is habitual. Life-size figures of this bone, viewed from the 



SHUFELDT.] 



OSTEOLOGY OF THE SPEOTYTO. 607 



three principal positions for the purpose of study and measurement, are 
offered to the reader in PI. I and PI. II, Figs. 5 and 6. The concave 
dorsal aspect of the body is smooth, being traversed in the median line 
by a very shallow groove that lies immediately over the base of the 
keel. This groove terminates, within five millimetres of the anterior 
border, in a little depression, at the bottom of which are discovered 
pneumatic foramina, two or more in number, leading to the anterior 
thickened vertical ridge of the carina beneath. Other minute openings 
for the admission of air into the interior of this bone are seen among 
some shallow depressions just within the costal borders. The bone does 
not seem to be as well supplied in this respect as it is in some other 
Owls. The costal borders supporting the transverse articular facets for 
articulation with the hoein apophyses occupy about one-third of the en- 
tire lateral border on either side anteriorly. At the bases of the major- 
ity of the depressions that occur between these facets are found other 
pneumatic foramina. The anterior border is smooth and rounded, with 
a median shallow concavity occupying its middle third. At its extrem- 
ities, laterally, the costal processes arise with a general forward tendency 
at first, but with their superior moieties directed backwards. The costal 
borders terminate at the posterior borders of the processes, at a higher 
level than the anterior sternal margin does at their anterior borders. 
The coracoid grooves are just below the anterior border. They are 
deep, continuous with each other, having a greater depth behind the 
manubrium in the median line than observed at any other point. Their 
general surface is smooth and polished, looking upwards and forwards, 
and lying principally in the horizontal plane. They melt away into the 
body of the bone laterally, at points opposite and not far distant from 
the posterior articulations on the costal borders. The margin that 
bounds them below is sharp, travels at right angles from the median 
line at first to a point posterior to the costal processes, then making a 
little dip downwards, then again curving upwards, disappears gradu- 
ally with the groove it bounds. That portion of it from the point where 
it changes its direction to its termination is described by authors as the 
subcostal ridge. The manubrium, occupying its usual position in the 
middle line, is comparatively small, quadrate in form, compressed below, 
slightly notched and flattened above, its posterior surface forming the 
inner anterior surface of the coracoidal groove. All the borders bound- 
ing the posterior parts of the bone are sharp ; the lateral one, taken 
from the apices of the costal processes to their other and lower termi- 
nations, are concave. As is the arrangement generally among Owls, 
the xiphoidal extremity of the sternum is four-notched, two on either 
side, the outer notches being the deeper. Both have rounded bases, and 
the processes that separate them are ample and possess rounded ex- 
tremities. The border upon which the keel ends posteriorly is square, 
though we have met with specimens in which it was slightly notched in 
the median line. The body is oblong, and, if we include the xiphoidal 
processes on either side, has a length half as long again as its width. 
The ventral and convex surface, like the dorsal, is smooth and presents 
but two points for examination. The pectoral ridge, faintly marked 
throughout its extent, originates on each side at a point near the outer 
borders of the coracoid grooves, running inwards and backwards, and 
dies away at the base of the keel near its middle. This little ridge 
denotes the line between the pectoralis major and minor. The keel is 
moderately well developed, the distance from the base of the manubrium 
to the carinal angle being equal to the distance from the same point at 
the base of the manubrium to the base of either costal process or outer 



608 GEOLOGICAL SURVEY OF THE TERRITORIES. 

anterior sternal angle. It is compressed, smooth, and thin, but its sta- 
bility is greatly aided by the carinal ridge on either side, which com- 
mences strong and well marked at the base of the manubrium, just 
within the anterior border running parallel with the latter, and disap- 
pears as it approaches the carinal angle. The anterior border of the 
keel is sharp and concave ; the inferior border is convex, with the edge 
slightly thickened. The point of intersection of these two borders an- 
teriorly is rounded and forms the carinal angle. The inferior border 
expands posteriorly, and the keel terminating a short distance before 
arriving at the posterior sternal border, the two become blended with 
the surface of the body of the bone. 

Sacral vertebrce ; pelvis; and coccygeal vertebra?. — In the sacrum of the 
Owl now under consideration, with the exception of a few faint lines in- 
dicating the original individuality of the vertebrae, these bones are 
thoroughly anchylosed together and to the ossa innominata. From in- 
spection of this compound bone in immature birds, we find the usual num- 
ber of sacral vertebrae composing the sacrum to be thirteen. The anterior 
face of the first possesses all the necessary elements for articulation with 
the last dorsal. The neural spine has a thickened crest that soon meets 
the ilia on either side ; its anterior edge is thin, and gives attachment be- 
low to the interspinous ligament. The neural canal is circular, and the 
prezygapophyses well marked. The articular facet of the centrum is in 
the vertical plane, with its curvatures similar to those ascribed to the an- 
terior facet on the centra of the dorsals. Theneurapophyses are broad 
and the diapophyses are strong and raised, with their enlarged ex- 
tremities expanded upon and firmly united with the iliac bones. There 
is but one pair of free sacral pleurapophyses 5 these are long and 
slender, articulating with the first vertebra in the usual manner, but 
the relation is much more intimate, as they touch the diapophyses for 
some little distance beyond the tubercula towards the capitula. The 
lower extremities of these ribs are terminated by little roundish knobs, 
which articulate with the hsemapophysis on either side, described as 
being inserted in the posterior border of the fifth sternal rib. View- 
ing the bone dorsal- wise, it is to be seen that the thickened crest of the 
neural spine of the first vertebra protrudes from the angle made by the 
ilia meeting it anteriorly to a greater or less distance. This broad and 
compressed crest, then continued backwards, is firmly wedged between 
the ilia until we pass the third vertebra ; at this point the ilia diverge from 
each other to another point just anterior to the acetabula, then converge, 
terminating in the posterior sacroiliac border within five or six milli- 
metres of each other. The sacrum completely fills in the lozenge-shaped 
space thus formed from the third vertebra — first, by continued broaden- 
ing and compression of the neural spine, that soon becomes one with the 
neurapophyses ; and, secondly, by the expanded extremities of the di- and 
par-apophyses, the processes themselves also taking due part. The in- 
tegrity of the surface is unbroken, save posteriorly, where a few pairs of 
foramina exist among the expanded transverse processes, increasing in 
size from before backwards. Anterior to a line joining the acetabula this 
surface is in the horizontal plane ; posterior to this line there is a decline, 
which declination is accepted also by the innominate bones ; this gives 
the entire pelvis a shape that seems to be characteristic of the majority of 
both the diurnal and nocturnal Baptores. The "ilio-neural" canals, here 
present, open by small apertures posteriorly, at about the point where 
the ilia commence to diverge, passing obliquely downwards and for- 
wards ; their anterior openings are large enough to allow a view of their 
internal walls. The neural spine that divides them throughout is com- 



shufeldt] OSTEOLOGY OF THE SPEOTYTO. 609 

pressed from side to side ; the ilia which form their outer boundaries are 
convex; the neuro-spinal crest forms the roof, the basal surface being 
deficient, formed merely bj the spine-like di- and par-apophyses of the 
vertebrae and the confluent neural arches. This first vertebra occupies 
the lowest level, the bird supposed to be standing as in PL I. Now, a 
line drawn mesial on the centra below, from the tirst centrum to the last 
gradually rises until opposite the anterior borders of the ischiadic fora- 
mina, then curves rather abruptly downwards to its termination. The 
centra of the first two or three vertebras are compressed from side to side 
to such an extent as to cause them to appear wedge-shaped, the common 
apex or edge being below; after that, however, they rapidly broaden, 
become compressed vertically and more cellular in structure; they are 
very broad from the fourth to the ninth, inclusive— then as rapidly be- 
come contracted as they approach the coccyx. Minute but numerous 
pneumatic foramina are seen at or near the usual localities. The largest 
foramina for the exit of the roots of any pair of sacral nerves is gen- 
erally in the fifth vertebra ; they decrease in size as they leave them 
either way. In the young, only the last few of these foramina are double; 
they are all double in the adult and placed one above another, a pair on 
the side of each centrum at their posterior borders, for the exit of the roots 
of the sacral nerves. The diapophyses of the anterior five sacral verte- 
bras are thrown out against the internal surfaces of the ilia, to which 
they are firmly attached, and act as braces to hold the engaged bones 
together. The parapophyses of the first, form facets for articulation 
with the sacral ribs; the second and third have none; in the fourth and 
fifth they also act as braces in the manner above described, joining the 
ilia just before their divergence commences. Reliance seems to have 
been placed entirely in the completeness of the sacro-iliac union in the 
last vertebras, for the apophysial struts terminate in that x>ortion of the 
pelvic vault formed by the sacrum itself, except in the last two verte- 
bras, where the parapophyses abut against the iliac borders. The para- 
pophyses of that vertebra which is opposite the acetabula are promi- 
nent, they being long and ample, reaching to the border and re-enforcing 
that part of the pelvis that requires it the most, the vicinity of the 
leverage for the pelvic limbs. In other Strigidce several apophyses are 
thrown out at this point. The posterior opening of the neural canal in 
the last sacral vertebra is subcircular, its diameters being about a milli- 
metre in length. This vertebra also possesses small postzygapophyses, 
looking upwards and outwards for articulation with the prezygapophyses 
of the first coccygeal vertebra; the articulating facet of the centrum is 
also small, long transversely, notched in the median line, the surface on 
either side being convex. At every point where the sacrum meets the 
iliac bones union is firm and complete, though both upon the internal 
and external surfaces the sutural traces are permanently apparent. 
The anterior iliac margins, as they diverge from the sacral spine, form an 
acute angle, concave forwards ; they have a well-marked rim or border, 
nearly a millimetre in width, raised above the general surface of the 
bone, which disappears on the outer borders as we follow them backwards. 
The two anterior and outer angles overhang the sacral and fifth or 
last dorsal pleurapophyses. From these last the marginal boundaries, 
which necessarily give the bones their form, are produced backwards and 
outwards to a point opposite the centrum of the third sacral vertebra, 
then backwards and inwards, forming at the above points two lateral 
angles. From the apices of the two lateral angles to where the borders 
terminate on either side in front of the acetabula with the pubic bones, 
the direction is such as to form a concavity on each side; the line joining 
39 H 



610 GEOLOGICAL SURVEY OF THE TERRITORIES. 

the bases of these concavities, points opposite the posterior openings of 
the ilio-neural canals, being the narrowest part of the pelvis. The upper 
and at the same time the inner margins of the bones in question, from 
the anterior and median angle, at first approach, soon to diverge from 
each other, andform the gluteal ridges and borders of those scale-like pro- 
jections of the posterior portion of the ilia that overhang the acetabula. 
Produced now as the "gluteal ridges,' 7 they tend almost directly back- 
wards, though very slightly inwards, to terminate in the ischial mar- 
gins. The preacetabular dorsal iliac surfaces are generally concave, 
while the postaeetabular, and at the same time that surface which 
occupies the higher plane, is flat, having a slope downwards and 
backwards, with a ventral reduplication after forming the rounded and 
concave posterior boundary of the pelvis. The preacetabular super- 
ficial iliac area is nearly double the extent of the postacetabular. 
The antitrochanterian facets that surmount the cotyloid cavities have 
the usual backward direction, though their surfaces look downwards, 
outwards, and a little forwards. The external surfaces of the ischia look 
upwards and outwards, having just the reverse direction ventrally. 
Posteriorly, these bones are produced beyond the ilia into finely pointed 
extremities, tending to approach each other. The slender pubic bones, 
after closing in the obdurator foramen on either side, touch and unite 
with the interior borders of the ischia as far as the pointed ends of the 
latter, beyond which they are produced nearly to meet behind. The in- 
terval between the free extremities of the pubic bones in some individ- 
uals, notably "birds of the year," is very slight, less than a millimetre 
sometimes, approaching a closed pelvis. The circular and thoroughly 
perforated acetabula are formed in the usual manner by the three pelvic 
bones. They have a diameter of about three millimetres, and their cir- 
cumferences are in the vertical plane. The ischiadic foramina are ellip- 
tical and large; they are, as usual, posterior to the acetabula and above 
the obdurator foramina. These last are also elliptical, and about one- 
third the size of the others. Should the major axes of these two ellipses 
be produced backwards, they would intersect and form an acute angle 
just within the posterior pelvic border. Viewing the pelvis ventral- 
wise, we observe, in addition to points mentioned when speaking of the 
sacrum, the reduplication of the ilia, forming pockets behind and inter- 
nally, that open outwards through the ischiadic foramina and inwards 
into the general pelvic cavity. The pelvic passage is subcircular, un- 
closed, with an average diameter of 1.7 centimetres vertically, and a 
little less transversely. The narrowest part of the pelvis measures 1.2 
centimetres, the widest 2 centimetres, being taken between the iliac pro- 
jections over the acetabula ; the average length, including anterior neural 
spine, is 3 centimetres. Pneumatic tbramina occur in the shallow an- 
iractuosities, between the antitrochanters and gluteal ridges in the ilia. 
None of the caudal vertebrce are grasped by the pelvis, the posterior ex- 
tremity of the sacrum always assisting to form the curve of the pelvic 
passage. The usual number of these vertebrae is seven, though occa- 
sionally an additional one is found, making eight iji some individuals. 
This enumeration does not include the modified and ultimate coccygeal 
vertebra, the pygostyle. They are all freely movable upon one another, 
and the first upon the last sacral vertebra* The articular facets upon 
the centra vary in shape throughout the series; that upon the first is 
long transversely, with a double convexity so arranged as to accommo- 
date itself to the one on the extremity of the sacrum ; they soon become 
uniform, to pass to the subcircular one existing between the last verte- 
bra and the pygostyle, on which it is concave. 



SHUEELDT. 



OSTEOLOGY OF THE SPEOTYTO. 611 



Tbe pleurapophyses and parapophyses are very rudimentary or en- 
tirely suppressed. Each vertebra bears a prominent neural spine, whicb, 
from tbe first to tbe sixth, inclusive, is bifurcated; in tbe last two it ap- 
pears as a mere primitive knobule. Tbe transverse processes are all 
deflected downwards and outwards, very small in tbe first and still more 
so in tbe last; are largest in tbe fiftb and sixtb. Prezygapopbyses are 
well marked; they reach forwards and articulate with tbe feebly devel- 
oped postzygapophyses. In a few of the posterior segments there ap- 
pears to be an effort on the part of tbe neurapophyses to overlap the 
vertebra next beyond them. The neural canal is pervious throughout, 
commencing in the first with a calibre equal to that in the end of tbe 
sacrum ; it gradually diminishes, and terminates in a minute, blind, con- 
ical socket in the pygostyle. Hypapopbyses are produced downwards 
in a few of the ultimate vertebrae. They hook forwards and articulate 
with the centrum of the vertebra next beyond them. Sometimes they 
are observed to be free, or rather resting upon a facette on the anterior 
margin of one centrum and extending over to the anterior margin of the 
centrum of the vertebra anterior to it, to meet a similar facette, as a 
tiny styliform process. The spinal column is completed posteriorly by 
the pygostyle — that ploughshare- shaped segment tbat articulates with 
the last coccygeal vertebra. Above its cup -shaped facet this bone arises 
as a laterally compressed plate, extending backwards and bifurcated at 
its extremity, as if to imitate the neural spines of the vertebrae of the 
series of which it is an ultimate appendage. Below the facet it projects 
forwards and completes the median sequence of hypapopbyses of the 
centra, being rather larger than any of them. The posterior curve is 
simply inflected downwards and forwards from its apex. 

The scapular arch — (See PI. I). 1 — The three elements that constitute 
this arch are all represented, and all independent or free bones; the 
coracoids articulate with the sternum and scapulae ; coracoids and clav- 
icle, connected by ligaments, lend their share to form or strengthen the 
shoulder -joints. Tbe coracoid, comparatively large and strong, forms 
in the usual manner an arthrodial joint of restricted movement with the 
sternum, its lower end being in the coracoid groove on the anterior part 
of that bone. The inner angle of its base is about 2 millimetres from 
the mesial line, and 4 millimetres intervening between it and its fellow 
of the opposite side in the groove. This extremity is broad, its outer 
angle being beneath the tbird sternal rib at its point of meeting the 
costal border; it is compressed from before backwards. The articular 
facet, looking downwards, backwards, and a little inwards, is trans- 
versely concave, with a slight dividing ridge, running antero-posteriorly, 
converting the general concavity into two smaller ones. The coracoid 
when in position is produced upwards, forwards, and outwards, making, 
with the vertical line through its base, rather an acute angle. A limited 
portion of the middle third of the bone only is subelliptical on section 
and at all shaft-like, due to the fact that the coracoid in this bird being 
perhaps less than the average length as compared with the size of the 
bird, and, secondly, to the unusually enlarged extremities, features ob- 
servable, more or less, in Raptores generally. The anterior groove of 
the upper extremity, that is arched over by the head of the clavicle 
above, is deep, and occupies fully the upper third of the bone. The co- 
raco-clavicular process springs, thin and compressed, from the inner 

1 It will be seen that in this figure, corresponding limbs, and other parts that are 
alike on either side of the body, have not been reproduced, it being thought the bet- 
ter way, as the bones on the side towards the observer would necessarily obscure the 
more remote ones, complicate the figure, and show nothing additional. 



612 GEOLOGICAL SURVEY OF THE TEEBITOEIES. 

side of the shaft of the bone, at junction of upper and middle thirds, to 
tarn upon itself, so as to be projected upwards, forwards, and a little 
outwards, terminating with an elliptical facet for articulation with the 
clavicle. The upper border of this process is concave lengthwise and 
articulates throughout its extent with the inferior margin of the acro- 
mial process of the scapula. The lower and thin edit of the eoraeo- 
clavicular process tends obliquely downwards, to be lost on the inner 
surface of the shaft of the bone near its middle. The outer wall of the 
anterior groove is formed by the coracoid itself, the process just de- 
scribed being really nothing more than a wing-like extension forming 
the inner boundary of the groove in this bird: it terminates above both 
clavicle and scapula in a rounded, tuberous head. Below this head, an- 
teriorly and still more inwardly, the coracoid affords a vertical, elou gated 
facet for the clavicle, while behind, looking a little outward, is the con- 
cave elliptical facet that constitutes about one-third of the glenoid cav- 
ity for the humerus, internal to which, and running first directly up- 
wards, then making a right angle and continuing forwards, a little 
upwards, and outwards, the last direction being the upper margin of 
the coraco-claviciilar process, is another facet, for the scapula. Behind 
and below, this bone displays one or two lines and depressions, bound- 
aries of muscular attachments. In the middle of the anterior groove. 
opposite the base of the coraco-clavicular process, the shaft of the bone 
is perforated : this r>erforation is elliptical lengthwise with the shaft, and 
passes directly through to make its appearance on the posterior cunvex 
surface just below the scapula. This foramen transmits a branch of 
that cervical nerve coming from between the twelfth and thirteenth cer- 
vical vertebra?. This nerve branch, after passing through the bone, is 
distributed to the under surface of the pectoralis minor muscle, and its 
filaments ascend among its fibres. This foramen is observable also in 
other Owls, as Bubo rirginianus. and in some of the diurnal E 
in Accipiter cooperi ; in very many birds it is absent. The scapula pre- 
sents little that is unusual in that bone among the class generally. It 
lends the additional two-thirds of articular surface to form the glenoid 
cavity with the coracoid: internal to this the acromion process extends 
forwards, touching the coracoid as described, and having a limited bear- 
ing on the clavicle. Posteriorly its blade-like length is produced, ex- 
panding, turning slightly outwards to terminate in an obliquely trun- 
cate extremity, with its point over the second dorso-pleorapophysial 
interspace. 

What the scapula lacks in interest is amply made up by the changes 
observed in the last bone of the group, the clavicle. This element is 
broad above, much compressed from side to side throughout : it spans 
the anterior groove of the coracoid and touches the scapula as described 
above, rapidly diminishing in size as it is produced downwards and in- 
wards by a gentle curve towards the fellow of the opposite side. The 
upper extremities in adult birds are separated by an average distance 
of 2.3 centimetres. If the sternum pointed to feebleness of flight in 
this little Owl. it is still further carried oat by the ill-developed clavicles. 
which constitute that arch in birds, where they are thoroughly and 
firmly united below, that assists to resist the pressure of the humeri when 
the wings are depressed in flight, and send them back to their former 
position after the completion of the action. In examining again PI. I. 
which represents the skeleton of an old male, we find this bone to be 
simply a pointed styliform process: in other individuals, and adults too. 
it does not even attain the length here shown : but. as if to bid defiance 
to all law or invariable* rule governing it. we again find in very young 



shufeldt.] OSTEOLOGY OF THE SPEOTYTO. 613 

birds cases where it becomes confluent with its fellow, forming a broad 
U-shaped arch, though never a very strong one. In a case of this kind 
the bone was finely cancellous throughout, with an extremely attenuated 
layer of compact tissue outside,. scarcely covering it. In PL I, and 
other individuals like it, the clavicles were pneumatic. Again, in both 
young and old, it may have any of its lower parts completed by carti- 
lage; it never displays a mesial expansion of bone at the point of 
confluence. We believe in the Barn Owl (Aluco) it anchyloses with 
the sternum at the carinal angle, by its bending backwards and meet- 
ing it at its lowest and median point. As already shown, the superior 
entrance of the anterior groove on the coracoid is a complete circuit, 
formed by the three bones of the group. The head of the coracoid 
overhangs it above; next below is the clavicle, closing it in anteriorly; 
lowest of all the scapula behind. A plane passed through the superior 
margins of this aperture would look up wards, inwards, and backwards. 
All the bones of the scapular arch are pneumatic, with the exception 
sometimes seen in the clavicle, and the foramina, to allow the air to enter 
their interiors, look into the inclosed groove of the coracoid just de- 
scribed. In the scapula the foramen is usually single and in the acro- 
mion process; single again in the clavicle, it is seen in the broadest part 
of the head, while in the coracoid there is generally a group of these 
little apertures, situated in the depression on the surface that overhangs 
this entrance to the coracoidal groove. 

As in many others of the family, in common, too, with not a few of the 
diurnal Raptor es, this Owl possesses, particularly the older individuals, 
an os humero scapular -e, of the usual form, that increases the articular 
surface of the shoulder-joint for the humerus. 

Of the upper extremity. — The upper extremity consists of ten distinct 
bones in the full-grown bird, omitting minute sesamoids that might ex- 
ist. These are the humerus of the arm, the radius and ulna of the fore- 
arm, two free carpals, the metacarpal, and four phalanges. (See PI. I.) 
The humerus is a long, extremely light and smooth bone, and when 
viewed from above in its position of rest, with the wing closed, it re- 
minds one of the curve in the small italic letter/, being concave above 
towards the scapula; and this bone is so twisted that this same curve 
is exhibited, though not quite as well marked, when viewing it laterally. 
The humerus is 5.5 centimetres long, subcylindrical on section at mid- 
shaft, at which point a minute aperture exists for the passage of the 
nutrient vessels that are distributed to the osseous tissue and its inter- 
nal lining. This foramen enters the bone very obliquely, its external 
orifice being nearest the proximal extremity. This end is well ex- 
panded and surmounted above by a strongly developed radial crest 
that overhangs the shaft slightly towards the palmar aspect. It occu- 
pies a line on the bone from the articular facet foi the shoulder-joint to 
an extent shown in PI. I. The ulnar crest, or lesser tuberosity, incloses 
quite an extensive fossa below, which acts also as a partial screen to the 
pneumatic foramina, for the humerus is highly pneumatic. They usu- 
ally consist of one circular opening, surrounded by a group of many 
smaller ones. In young birds a very large foramen is generally pres- 
ent; this closes in as age advances. Between the two tuberosities is 
the vertical and elliptical convex facet for articulation with the glenoid 
cavity of the shoulder-joint, constituting the "head of the humerus." 
The radial crest displa\ s palmad, a ridge for the insertion of the tendon 
of the pectoralis major. The distal end of the humerus is also ex- 
panded in the vertical plane and gentlj 7 convex anconad, the reverse 



614 GEOLOGICAL SURVEY OF THE TERRITORIES. 

condition of the proximal extremity. It presents, for examination, the 
articular facets for the ginglyrnoid joint it forms with radios and nlna, 
and the superior and inferior condyles. The larger, and at the same 
time the superior, of these two facets is intended for the cup-shaped 
depression in the head of the radius, as well as a portion of the articu- 
lar surface on the ulna. It is ovoid in form and placed obliquely on the 
bone, the inferior end of the long axis of the oval being situated the 
nearer the proximal extremity of the shaft. This facet is separated 
from the trochlea surface for the ulna by a well-marked depression : this 
latter is a knob-like tubercle when compared with the radial facet. 
The condyles and the entire articular surface are about in the same 
plane posteriorly ; that is, neither increases the length of the bone, 
one more than another. Passing from the trochlear surface for the 
ulna towards the inner aspect of the shaft, there is to be observed 
a shallow depression, which corresponds to the olecranon fossa of 
human osteology, and in full extension of the limb allows room for 
that process of the ulna in this bird. The radius has an average length 
of 6.6 centimetres, and the ulna a corresponding length of 6.S centi- 
metres, so that their distal extremities, when articulated, as we exam- 
ine them in the closed wing, extend beyond the head of the humerus. 
In this position also the radius occupies a higher level than the ulna, 
and is the innermost bone of the two. The radius is slender, the trans- 
verse diameters of its subcylindrical shaft varying but little throughout 
its extent, though its extremities are expanded. From the elbow-joint, 
when the two bones are in position, it at first diverges from the ulna at 
a moderate curve, to approach that bone again by a more gentle incli- 
nation to nearly absolute contact at the junction of middle and distal 
thirds ; from this latter point it lies parallel with the ulna to the wrist 
The head of the radius is elliptical, being crowned by a depression for 
articulation with the oblique facet on the distal end of the humerus. 
Beyond, below, and to the outer aspect of this facet is another of similar 
form, though convex for articulation with the ulna, while still more ad- 
vanced toward the distal end we find the bicipital tuberosity, and still 
more distally, the minute nutrient foramen ; all of the bones beyond the 
humerus being non-pneumatic. The distal extremity of the bone in 
question is terminated by a little fan-like expansion that caps the ulna 
and articulates by its anterior convex margin with the scapho-lunar of 
the wrist. It is marked above by the longitudinal groove for the tendon 
of the extensor metacarpi radialis longus. The shaft of the ulna is 
nearly three times as large as that of the radius. Its outer 1ml: > 
straight, its inner curved towards the humerus, thereby increasing, at 
the proximal moiety, the interosseous space, by the assistance of the 
opposite curve made by the radius. The stronger end is the one in- 
volved in the formation of the elbow-joint : here is to be observed 
the depression for the head of the radius, or the lesser sigmoid cavity, 
while the articular surface beyond that occupies the entire end of the 
bone, directed downwards, iuwards, and backwards, presents for exam- 
ination the greater sigmoid cavity, the olecranon and coranoid pro esses, 
and the cavity for articulation with the oblique facet of the humerus. 
The greater sigmoid cavity is subcircular and of some depth ; its lower 
aud produced lip represents the coranoid process, as does its upper, 
better marked, and more tuberous prolongation represent the olecranon 
of andranatomia. Extending radiad is another concave, quadrate, ar- 
ticular facet for the oblique tubercle of the humerus, as the first-men- 
tioned concavity articulate* with the ulnar tubercle or trochlea. A 
little beyond this articular surface are various small tuberosities and 



shufeldt] OSTEOLOGY OF THE SPEOTYTO. 615 

depressions for the origin and insertion of muscles. Approaching the 
wrist, the shaft is seen to be generally smooth, and diminishes in cali- 
bre at junction ot middle and proximal thirds, in the locality of the 
nutrient foramen, while along its entire length, at certain intervals, are 
the slight elevations for the apices of the quills of the secondaries. 
The distal extremity of the ulna enters into the formation of the wrist- 
joint ; it is not nearly as large as the proximal end. The articulating 
surface has a deep mesial cleft in the vertical direction, limited exter- 
nally by an elliptical curve, internally by a double, tuberous kuob for 
articulation with the irregularly formed cuneiform of the carpus, while 
above is a roughened surface that is covered by the expanded end of the 
radius. 1 

The carpus is composed of the scapho-lunar, os magnum, and cuneiform. 
The scapho-lunar articulates with radius, os magnum, and ulna. The 
radial articulation is a rather deep and elliptical concave facet, its lower 
border gliding over the ulna, w r hile the distal end of the radius plays in 
the concavity. The opposite face of this six-sided little bone is also 
smooth, and is a nearly flattened surface that articulates with os mag- 
num. The upper and lower surfaces, as well as the ends, are simply 
roughened and fashioned to give the proper form to that part of the joint 
into which it enters, and tor the attachment of ligaments. Os mag- 
num has become confluent with the mid-metacarpal, forming its troch- 
lear surface for articulation with scapho-lunar, cuneiform, and ulna. The 
cuneiform is an extremely irregularly shaped bone; it appears to be 
rather the larger of the two free carpals, and is the lower in regard to 
position. It articulates with ulna and os magnum, simply. Its outer 
ulnar facet is elliptical and shallow, monopolizing the entire face of the 
bone; its inner facet is very irregular, being formed so as to accommo- 
date itself to the ulnar tubercles, with which it articulates. Projecting 
towards the metacarpus, this little bone has two prongs or limbs, the 
inner aspect of the extremities of each possessing a subcircular facette 
that articulates, the outer and shorter limb with the internal trochlear 
margin of os magnum, on the same side ; the inner and longer limb strad- 
dles the metacarpal and glides over the surface, during movements of the 
joint, at a point about w 7 here magnum becomes confluent with mid-meta- 
carpal. The cuneiform has also attached to it ligaments that enclose 
the wrist-joint beneath — capsular ligaments of the carpus. 

The metacarpus is formed in the usual manner, by the amalgamation of 
the index, medius, and annularis metacarpals, the first, second, and 
third, respectively. It is 3.3 centimetres long, articulating with scapho- 
lunar, ulna, and cuneiform at its proximal extremity by means of os mag- 
num, that has become anchylosed with mid-metacarpal and the pha- 
lli the October number of the Bulletin of the Nuttall Ornithological Club, for 1881, 
Cambridge, Mass., I published an article entitled "On the ossicle of the Antibrachium 
as found in some of the North American Falconidse"; wherein I described a very inter- 
esting sesamoid that was found above the carpal articulation, on the radius, iu a speci- 
men of Circus hudsonius. This sesamoid I named the os prominens, but subsequently 
ascertained that it had been noticed by Milne-Edwards, and after this naturalist, by 
Mivart (Lessons in Elem. Anaf., p. 320, Lond., 1877). My attention was afterwards 
called to its occurrence in the Owls, by Mr. Forbes, prosector to the Zoological So- 
ciety of London, and again by Mr. F. A. Lucas, of Rochester, New York, the latter 
observer finding it in Bubo and Nyctea. Subsequently I found it in other American 
Owls, and have no reason to doubt but that it will be found in Speotylo, though I have 
not material at hand, now, to confirm it. Additional literature and remarks upon 
this sesamoid will be found, in following numbers of the Bulletin quoted above; one 
in the January number, 1882, by Mr. J. A. Jeffries. It does not sbow in the carpus of 
Bubo virginianus (Plate III, Fig. 11), accompanying this article, because at the tiii& 
I drew the figure I was not aware of its occurrence in that Owl, or the joint repre- 
sented would have been placed so as to exhibit it. 



616 GEOLOGICAL SURVEY OF THE TERRITORIES. 

langes at its distal end. The first metacarpal is short, and fused with 
the second jnst anterior to the boundary of the trochlear surface of os 
magnum: it makes an angle with the shaft of the second metacarpal, 
its extremity being directed upwards. At its base, close to The shaft of 
mid-metacarpal. it bears a uniform facette for articulation with the index 
phalanx, a free, three-sided, pointed little bone, about 9 millimetres in 
length. The second metacarpal is straight : its enlarged proximal ex- 
tremity is formed chiefy by the confluent os magnum: its shaft is in- 
clined to be subtrihedral. with its broadest face looking forwards: its 
distal extremity is terminated by a knot shaped enlargement, that is 
still further enhanced by the continence with the third metacarpal. It- 
bears a digit composed of two phalanges, the proximal one bearing on its 
posterior border, for nearly its entire length, a quadrangular expansion, 
that has a raised margin, leaving a single concavity radiad : a ■ innlar con- 
cavity occurs on the ulnar side, but is there divided by a ridge, s oping 
downward into two shallow depressions. This little b me sun 'what re- 
minds one of a cleaver, with the end of its handle attached to the meta- 
carpus. It supports at its distal extremity the second phalanx ot this 
digit, a bone having very much the same appearance and shape as the 
index digit. 1 only being longer and more pointed. The proximal ends 
of all the phalangeal segments are more or less expanded, in order to 
support the ample facets of articulation that occur among them, and 
the metacarpus. The third metacarpal is expanded transversely above, 
slender below, where it falls a little beyond the inedius after its conflu- 
ence with it. It also has a small, pointed phalanx, freely attached to 
its distal extremity, and lying in that recess formed by the shaft and 
posterior expansion of the first phalanx of the second digit. At a very 
early date, comparatively, in the life of this Owl. ossification is normally 
extended to many of the tendons of important muscles of the antibra- 
chium and pinion. 

Of the pelvic limb. — The lower extremity is composed of twenty dis- 
tinct segments, including the patella, or just double the number found 
in the pectoral limb. This increase will not surprise us when we recol- 
lect the greater number of small bones devoted to the foot above th 
found in the hand. Its most striking feature, next to those osteological 
characteristics common to the family, is its extreme length, due princi- 
pally to the tibia and tarso -metatarsus. All the bones of the lower 
limb in this species are non pneumatic. The femur is comparatively of 
good size and strong : articulated in the usual manner, it measures I 
centimetres in length and 7 millimetres across the condyles at their 
widest | art. At the proximal extremity, externally, above the shatt. 
there is a flat and roughened surface, bounded above by the curved 
trochauterian ridge. This surface forms the major part of the great 
trochanter. There is no trochanter minor present. The trochauterian 
ridge is the highest part of the bone, when it is held vertically : it lies 
in the antere-posterior plane, with the femur in its natural position, the 
bird standing erect : from it. sloping directly inwards and occupying the 
remainder of the summit between it and the head, is a smooth articu- 
lar facet, broadest externally, merging into the globular head internally. 

l Mi. J. A Jeffries, in a very interesting article entitled " On tlie Fingers of Birds'' 
(Bull. Xutt. Ornith. Club, January. 1881, p. 6). endeavors to settle the argument 
upon the homology of this joint. This author says: "Whether the metacarpus are 
the I— III or the II-IV has been a mooted question. Eolleston. Huxley, and Gegea- 
baur holding the first view, and Owen. Wyman. Morse, and Cones holding the second 
view."' I have always maintained the view expressed by the latter gentlemen men 
tioned: the fact, however, that the first phalanx of manus in Ave* is tlie homologue of 
poliex of the pentadactyle limb seems to be gaining ground. 



shufeldt.] OSTEOLOGY OF THE SPEOTYTO. 617 

With the head it constitutes the articular surface for the pelvis — it be- 
ing opposed to the antitrochanterian facet of the ilium, while the caput 
temoris plays iu the cotyloid ring. The excavation for the ligamentum 
teres on the latter is conical and deep, consuming a good part of the 
bone; it is situated on its upper and inner aspect. In looking into the 
relation existing among head, neck, and shaft of the femur of this bird, 
we must observe that if the straight line lying in the middle of the 
surface of the internal aspect of the shaft were produced upwards, it 
would pass through the centre of the facet at the summit — if anything, 
nearer the trochanterian ridge than it does to the head. This facet also 
is notably narrower just before arriving at the head than at any other 
point. Again, the plane passing through the external and circular bound- 
ary of the head makes an angle of a good 45° with this line, so that 
with these facts in view we can hardly assert in the case of the species 
before us, as do some authors on comparative anatomy in describing 
this bone in general, that the head of the femur is either nearly at 
right angles with or is sessile with the shaft. It would appear, though, 
that it has quite as much of a neck to boast of as the anatomical neck 
of humerus or the neck of the scapula in works on human anatomy. 
The shaft throughout its length, until it begins to approach the distal 
condyles, where it is subcompressed and expanded antero-posteriorly, 
is nearly cylindrical, bent slightly backwards at its lower end, ami 
offers for examination merely the intermuscular ridges, with the liuea 
aspera, feebly marked, and the nutrient foramen, all of which maintain 
their usual positions on the bone. At the distal extremity the rotular 
canal, the intercondyloid notch, and the popliteal fossa are all strongly 
produced, giving due prominence to the condyles, internal and external, 
between which they form the dividing tract. The external and lower 
condyle is divided in two by a vertical excavation, deepest above. Of 
the two facets thus formed, the inner articulates with the tibia, the 
outer with the head of the fibula. The external surface of this condyle 
is flat and continuous with the shaft. The inner condyle, broad poste- 
riorly, has a slight depression in the surface that bounds it on the tibial, 
side, and as a rule the usual sites for ligamentous attachments about 
this extremity are at best but feebly represented. The patella, encased 
in the tendon of the quadriceps femoris, is situated about 3 millimetres 
above the rotular crest of the tibia, anteriorly, having the form of au 
oblate hemispheroid with its base directed upwards, the long diameter 
of which measures 3.5 millimetres. The tibia is the longest bone in this 
bird's skeleton, and at the same time ? taking this length into considera- 
tion, the least curved or bent along the shaft; it has, however, a slight 
and just appreciable gradual curvature forwards that is most apparent 
about the junction of middle and upper thirds. Its average length, 
measured on the inside, is 6.7 centimetres ; its extremities being ex- 
panded for articulation, above with the femur, below with the tarso- 
metatarsus. These expansions are of about equal dimensions, though 
differing vastly in form, in this respect being unlike some of the diurnal 
Baptores, in which the distal condyles constitute the smaller end of the 
bone. 

Among the most important points presented for examination about 
the head is the articular surface that crowns it above for the condyles 
of the femur. This is subquadrate in form, uneven, highest at the in- 
ner and anterior angle, sloping gradually to the opposite one, bounded 
almost entirely around by a raised margin, that is most feebly devel- 
oped posteriorly, and at a point anterior to the head of the fibula, where 
it is absent. In front this border may be nominated the rotular or epi- 



618 GEOLOGICAL SURVEY OF THE TERRITORIES. 

enemial ridge, though it is no more prominent there than at any other 
point, but in many birds it is so produced as to form a process of 
soine size, to which these terms are applied. Externally and poste- 
riorly the margin is roughened for the attachment of ligaments that 
bind the head of the diminutive fibula to this bone. In the middle of 
this articular surface is to be seen a tuberosity, on either side of which 
are the depressions for the femoral condyles. Produced downwards, 
anteriorly from the rotular ridge are the cnemial ridges; these have their 
crests bent slightly outwards, and they merge into the shaft below, 
abreast the superior point of the fibular ridge. Of the two, the outer 
or ecto-cnemial is the shorter ; that is, it does not extend so far down 
the shaft as the inner or pro-cnemial. They have between them an 
ovate concavity, with the larger end above, the lower end subsiding 
upon the shaft with the ridges themselves. The vertical elevation on 
the external aspect of the shaft for articulation with the fibula runs 
down the side but a short distance; a little below its abrupt termination 
may be observed in a line with it, the nutrient foramen, entering very 
obliquely from above downwards. After leaving the fibular ridge as far 
as the point where the bone begins to expand transversely at the distal 
extremity, the shaft is remarkably smooth and nearly cylindrical. This 
transverse and distal expansion is checked, both anteriorly and pos- 
teriorly, by abruptly meeting the distal condyles, the point of meeting 
perhaps being rather the higher behind. The condyles, differing but 
little in size, are singularly uniform as to shape, with their curved sur- 
faces downwards, being flat on their outer aspects, with a raised rim 
bounding them in each case. They stand out prominent and apart. 
Anteriorly their convex surfaces are the widest, behind they slightly 
approach each other, and the articular convex surface is narrowest on 
the outer condyle. The intercondyloid notch is deep, and appears 
equally well marked throughout its extent. Immediately above it, ante- 
riorly, there is a deep triangular depression ; another, and more shallow 
one, is found behind in the corresponding locality. Up the shaft a short 
distance on the inner side, anterioily, is a little tubercle, to which is at- 
tached the ligament that binds down some of the strong tendons of the 
extensors. This ligament crosses the anterior triangular depression 
mentioned above, obliquely, to be inserted near the external condyle 
superiorly. This is the arrangement also in Bubo virginianus, but in 
some of the Hawks this ligamentous bridge has become thoroughly ossi- 
fied, forming a strong bony band across the concavity in question. It 
is interesting to remark here, however general the rule may be as ap- 
plying to the diurnal and nocturnal Raptores, that whereas this band is 
ligamentous in the tibia in some of the Owls, a bony one fulfilling the 
same function is found in them just below the head of the tarso meta- 
tarsus ; these conditions are just reversed among some of the Hawks. 
In fact, we know of no exception to the arrangement just mentioned for 
the Owls; even Sumia funerea has this bony bridge on the tarso-meta- 
tarsus very prominent, and on the inner side of the upper third it being 
much in the same position as we find it in the well-known exception to 
the Hawks, that is, in Pandion. In short, among the Raptores it seems 
to be found among those birds that possess the reversible toe. Usu- 
ally, in old birds of this species, the fibula is firmly anchylosed to the 
entire length of the fibular ridge of the tibia ; arching outwards, its head, 
surmounted by an antero-posteriorly elongated iacet, rises a little above 
that bone at the point where it is attached to it by ligament. This is 
the larger part of the shaft in regard to size. Below the ridge this bone 
becomes simply a delicate little spine, that emerges into the shaft of the 



shufeldt] OSTEOLOGY OF THE SPEOTYTO. 619 

tibia at about the junction of middle and distal thirds, though it may 
be traced after this as far as the middle of the outer condyle, where it 
terminates by a minute tubercle. The head is notched externally, near 
the centre, and has lodged at that point a small sesamoid that is in the 
lateral ligament of the knee-joint. Posteriorly on the shaft, about midway 
down the superior tibiofibular anchylosis, we observe a small tubercle 
for the insertion of the tendon of the biceps. The long segment that 
exists between the tibia and the phalanges of the pelvic limb is the bone 
tarso-metatarsus, or the confluent metatarsals of the second, third, and 
fourth toes with certain tarsal bones at its proximal extremity. It meas- 
ures down the anterior aspect, mesially, 4.6 centimetres, and has its ex- 
tremities enlarged for articular purposes,in common with other long bones 
of the skeleton. At its proximal end the bone presents superiorly two con- 
cave articular surfaces for the condyles of the tibia. They appear nearly 
on a level with each other, the bone being held vertically. The inner and 
larger of the two is elliptical in outline, antero-posteriorly ; the outer and 
smaller is lashioned off behind by a tuberous process, directed upwards 
and outwards. Between these two surfaces arises a prominent tuber- 
osity, that in the articulated limb enters the intercondyloid notch of the 
tibia quite accurately, and is intended for a ligamentous attachment. An- 
teriorly and internally a groove exists that runs down the shaft, to dis- 
appear a little above its middle. This canal is deepest immediately be- 
low the articular expansion, and is here bridged over by a little arch of 
bone, a millimetre in width, that serves to bind down and hold in its 
proper place the tendon of the long extensor of the toes. Posteriorly 
there is a much deeper and longer tendinal canal, that extends the entire 
length of the shaft, being shallowest at the middle and most capacious 
at the proximal extremity ; this is bounded above and internally for a 
short distance below the head of the bone by the calcaneal process, a 
thin lamina of bone that has a foramen near its base ; this process is 
surmounted by an elliptical and compressed tuberosity, placed vertically. 
The opposite wall, above, of this groove is also thin, and extends, in 
common with the calcaneal process, directly backwards. There are two 
other foramina seen at this end of the tarso-matatarsus ; one just at the 
external termination of the bony bridge mentioned above, and the other 
outside and a little above it. Their posterior openings are immediately 
behind the anterior ones, or, in other words, they do not pierce the shaft 
in any way obliquely. The shaft of this bone is nearly square on sec- 
tion for the major part of its extent, being encroached upon, however, 
both before and behind, by the aforesaid tendinal grooves. The tendons, 
especially those that occupy the posterior canal, are very prone to ossi- 
fication, forming quite sizable bones in the adult, the largest of these 
being equal to the fibula in bulk, exclusive of course of the head of that 
bone, and not being as long. Returning to the tarso-metatarsus, we find 
at its distal extremity, for examination, the trochlear that articulate 
with the rear segment of all the toes except hallux. Viewing this end 
with the bases of these trochlese towards one, we find the general out- 
line made by them to be crescentic, with the horns having a tendency 
to approach each other behind. The outer trochlea is the highest and 
longest from before backwards; the other two are anout on the same 
level, the inner one having a posterior and internal process, while the 
middle one is possessed of a median cleffc traversing its face antero- 
posterior^-. They are sharply divided from each other by narrow slits, 
that extend up as far as the articulating part, and are continued on the 
anterior aspect of the shaft for a short way as delicate groovelets. A 
foramen is situated in the outer of these, that gives passage to the ante- 



620 GEOLOGICAL SURVEY OF THE TERRITORIES. 

rior tibial artery, and is comparatively larger than usually seen in the 
Owls. Behind, the tendinal groove expands, and is bounded distally by 
the concave border formed by the trochlea. Upon its internal margin, 
just above the extremity of the bone, it shows an elongated but feebly 
marked depression of about 3 millimetres in length. This facet articu- 
lates with the os metatarsale accessorium, which is joined to the bone 
by ligament. This little bone in this bird has an average length of 4 
millimetres. It is twisted upon itself, and bears upon one border a con- 
vex, smooth surface for the tarso-metatarsus, while distally it has an 
articulating surface, resembling more the mid-trochlea than any other, 
for the proximal segment of the hallux. Above it is sharply grooved 
for the tendon that goes to that toe. The toes are four in number, and 
their bony segments follow the rule that governs the greater part of the 
Class Aves; that is, first, second, third, and fourth toes have 2, 3, 4, and 
5 phalanges allotted to them, respectively. The first phalanx of the 
hind toe is more compressed from side to side than in the other toes, 
possessing more of the characteristics of the second joints. Its posterior 
facet, that articulates with the accessory metatarsal, fits accurately into 
the cleft surface seen on that little bone. Anteriorly the facet has a 
median groove, forming two vertical convexities for the double concave 
facet on the claw, with its dividing ridge. The claws are all a good deal 
alike, varying in size, the rear one being the most compressed laterally. 
They are pointed, arched, and nearly conical, the horny theca? that cover 
them during life only being grooved on the under side. Their proximal 
ends have an articulating facet for the next phalanx behind them ; this 
is so arranged that they can be more smartly flexed than any of the 
other joints of the foot, due to the convex articulating surface extending 
well beneath on the phalanx they meet. On the under sides of their 
proximal extremities is a tuberosity for the attachment of the flexor 
tendons ; it has on either side, below, an oval foramen to allow vessels 
and a nervelet to pass to the extremities of these ungual phalanges. The 
first joint of the second toe, and the first and second of the third, are 
thickest and short, articulating internally with the tarso-metatarsus, and 
having their facets so arranged as to allow" of motion only in the one 
plane. These bones may almost be said to interlock with each other, 
with their superior projecting processes behind fitting closely into the 
deep groove intended to receive them on the anterior faces of the joints 
to their immediate rear. The other underscribed phalanges of these two 
toes resemble the proximal segment of hallux. The fourth or outside 
toe possesses five phalanges, but the three innermost segments are very 
short, and are really nothing more than one of the middle type of pha- 
langeal bones, such as the third on the mid-toe, divided into three nearly 
equal parts, the proximal and distal pieces retaining all the character- 
istics of that bone, while the middle segment is simply a mid-section of 
the shaft. This arrangement, however, together with the manner in 
which the proximal phalanx, if it may be termed so in this bird, articu- 
lates with the long and elevated trochlea on the tarso-metatarsus, gives 
this toe a versatility and a power to be thrown outward and, to a limited 
extent, to the rear, not enjoyed by any of the other toes, constituting one 
of the most interesting anatomical features that we find in the family 
Strigidw. 



622 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE I. 

The skeleton of Spedtyto cunicularia hypogosa. 

Note. — The drawings from which the lithographic plates and the woodcuts of the 
hones in these monographs were made were executed hy the author ; all the cuts of 
external characters and illustrations of the hirds are from Baird, Brewer and Ridg- 
way's North American Birds, and kindly lent me hy Professor Baird. 



624 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE II. 

The skull, sternum, pelvis, etc., natural size. 
Fig. 1. The skull from above. 
Fig. 2. The skull from below. 
Fig. 3. The mandible from above. 
Fig. 4. The pelvis from below. 

Fig. 5. Anterior view of sternum and first dorsal vertebra, with its corresponding 
vertebral and sternal ribs. 
Fig. 6. The sternum from below. 



626 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE III. 

Various bones of the skeleton. 

Fig. 7. Anterior view of skull, the lower mandible having been removed. 

Fig. 8. H, humerus; i, ulnar tubercle ; i', oblique tubercle for radius and ulna; Ji, 
radial crest ; S A and S' A', scapular arch ; S and S', scapula ; C and C, clavicle ; Cr 
and Cr', coracoid ; /, perforating foramen. 

Fig. 9. R', humerus; h", radial crest; pf, pneumatic foramina; S A" and S A'", 
scapular arch ; S" and S'" } scapula; C" and C", clavicle; Cr" and Cr'", coracoid ; 
/', perforating foramen. 

Fig. 10. H A, hyoid arch ; a, superior view of atlas; a', the same viewed laterally ; 
&, the axis ; o, its odontoid process. 

Fig. 11. Right carpus of Bubo virginiamis, outer aspect, with the bones composing 
it moved partly from their normal positions to show articulating surfaces; rd, radius; 
ul, ulna; s, scaphoid; c, cuneiform; ra, metacarpus; d, index digit. 

Fig. 12. Right radius and ulna, Speotyto, inner aspect ; u, ulna ; r, radius ; y, articu- 
lar facet for oblique tubercle of humerus ; y' for ulnar tubercle of humerus. 

Fig. 13. The same bones, inferior surface, when in position and the wing closed ; 
r', radius; n', ulna. 

Fig. 14. Posterior surface, right metacarpus. The differences in form and position 
of such portion of the articular surface in the metacarpus as is shown by z and z', in 
Figs 11 and 14, between Bubo and Speotyto, are here seen; flat and rounded below in 
the first, prominent and pointed in the second. 

Fig. 15. Anterior surfaces, right tibia and fibula ; F, fibula ; T, tibia. 

Fig. 16. Posterior surfaces, same bones; F', fibula; T, tibia. 

Fig. 17. Right femur; v, posterior surface ; v', anterior surface. 

Fig. 18. Anterior surface, right tarso-metatarsus ; m, bony bridge over tendons ; j, 
foramen for anterior tibial artery ; e, facet for outer toe, I, for middle, and t for inner 
toe. 

Fig. 19. Posterior surface same bone : f, the foramen for the anterior tibial artery ; 
e" facet for outer toe, I" for middle, and t" for inner toe; Jc, facet for os metatarsale 
accessorium. 

Fig. 20. A, right os metatarsale accessorium, superior surface ; B, base or inferior 
surface of right tarso-metatarsus ; e', facet for outer toe,*', for middle, and t' for inner 
toe. The section of the shaft shows just above the middle facet, on the posterior as- 
pect, ranging near the middle third of the bone. 



OSTEOLOGY OF EREMOPHILA ALPESTRIS. 



By E. W. Shufeldt, M. D. 

Captain, Medical Department, United States Army. 



The lltli of March, 1880, was a particularly severe day at Fort Fetter- 
man 1 A violent wind and snow storm prevailed during the entire 
twenty-four hours. In the creek bottom, below the fort, where the wind 
had exposed the ground of some land that had been used for gardening 
purposes the year before, thousands of Horned Larks congregated. 
They seemed disinclined to vacate their partially sheltered position, pre- 
ferring to face the few death dealing fires I delivered them rather than 
be tossed over the prairie by the freezing storm. At each shot, the flocks 
arose, skimmed low over the ground, soon to alight a^ain. These sim- 
ple manoeuvres afforded me abundant opportunity to secure many speci- 
mens, and several hundred were taken. As they afterwards lay upon 
the table in my study, one would almost have said, before submitting 
them to careful scrutiny and examination, that not only was true alpes- 
tris represented, but both the varieties, leucolcema and chrysolcema, de- 
scribed by modern writers. Certainly it was that there were many 
shades of their normal coloring among them, accompanied by differ- 
ences in size that were not due to sex. I have never seen the black 
pectoral crescent of this bird in the low position in which Audubon rep- 
resents it in his work (B. Am., Vol. VIII, pi. 497), where he figures his 
Alauda rufa, the Western Shore Lark. 

Before proceeding to the study of the skeleton of this interesting 
species, we will remind the reader that of all the genera that go to make 
up the family A laudidce, or Larks, but few species have fallen to the lot 
of our North American avifauna. We have the Sky Lark, Alauda ar- 
vensis, 2 and the subject of the present monograph, with its two varieties, 
E. alpestris leucolwma and chrysolwma, mentioned above. 

The Skull — (PI. IV, Figs. 22, 25, and 26). — It is a striking characteristic 
in the skulls of nearly all adult birds that certain bones become firmly 
united, their sutures entirely disappearing ; perhaps in no species ot the 
highly organized suborder Oscines has this almost universal avian feature 
been so thoroughly carried out as in our present subject, the Horned 
Lark. Occasionally we do find, however, a trace to guide us'in locating 
the original boundaries of the primitive elements, even among the Os- 
cines, as the sutures, amidst the parietals and frontals in the cranium of 
Lanius, when maceration is carried to a high degree, but in Eremophila, 

1 Wyoming Territory, United States, lat. 42° 23' 35" N., long. 105° 21' 4" W. 

2 Alauda arvensis, Linn. — Cf. Dresser & Sharpe, Birds Enr., pt. — , and B. B. & R. 
Hist. N.Am. B.,ii. 1874, 136 (Greenland and Bermuda), and Bull. U. S. Nat. Mus., No. 
21, p. 21)3, and Coues' Cheek-list of N. A. B., 2nd ed., 1882, p. 33, No. 85. 

627 



623 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



as already stated, there is a total absence of any such indication. If we 
remove the lower mandible from the skull in any of the Class Aves, and 
place the remainder on the horizontal plane, with the basi-cranii down- 
wards, we observe that in different skulls there exists in this position 
differences in equilibrium, and differences in, what we will call, the an 
terior and posterior bearing points, or the points upon which this part 
of the skull rests upon the horizontal plane. To illustrate this in the 
skull we are studying, we find, when placed as directed above, that 
its equilibrium is quite stable, and that it rests posteriorly upon the tym- 
panies, anteriorly upon the tip of the superior mandible, which consti- 
tute, respectively, its posterior and anterior bearing points. In this 
case there is but one anterior bearing point, with two posterior ones. 
This is a very common result, but there are at the same time many ex- 
ceptions to it, as in Numenins, and many species of the family Anatidw. 

Again, if we erect a perpendicular from one of the posterior bearing 
points, or the posterior bearing point, fur sometimes it is the condyle, 
we find that the planes passing through the circumference of the fora- 
men magnum and the occipital vertebra, and the point where the foot 
of this perpendicular and the posterior bearing points coincide, make 
certain angles with the horizontal plane (the ordinary horn protractor 
is the best instrument to take these angles with), which we wi*l call, re- 
spectively, the angle of the foramen and the angle of the base. These two 
angles, in many instances, practical ly coincide, as in our Lark, where 
they make an angle of 40° with the horizontal plane. In the cut, H H' 
is the horizontal ylane j a the anterior and j? the posterior bearing points. 

These angles also differ in many birds ; e. #., the anterior bearing 
point in Ardea herodias is the tip of the upper mandible, the posterior 
ones being the inner of the three facets ou each tympanic; the angles 
of the planes of the base and foramen about coincide, and is 50°. In 
many of the Owls and diurnal birds of prey, the bearing points being 
the same as in the last example (it being, however, the inner facet of 
two on the tympanies, as a rule), the combined augles, or either of them 
separately, is very small, or the base and foramen may be found to lie 
nearly in a plane parallel to the plane upon which the skull rests, or the 



9fi° 




angles are 0°. 



We see then that in the present case, the bearing points 
being given, the angle of the combined planes is 40°, which fact, even 
without actually taking the angles in question, conveys to our minds 
about the " pitch" or relation of thebasi cranii to the other salient feat- 
ures of the skull. Taken accurately, these angles, it is obvious, would 



shufeldt.] OSTEOLOGY OF THE EREMOPHILA. 629 

have a certain value when we come to compare the various skulls of the 
Class. 

The primary elements of the occipital, or first cranial vertebra, have 
become completely fused together, and with such other bony elements of 
the vertebra beyond, of the mesencephalic arch, with which they usually 
articulate. The well-marked superior curved line that limits muscular 
attachment above would seem to be, and in all probability is, about the 
position of the lambdoid suture, and the superior boundary of the bone 
we are describing. This curved line descends and is gradually lost along 
the boundaries of the mastoids and occipitals on either side. Externally 
and inferiorly we find the occipital pierced by the usual foramina of the 
basi-cranii. The group for the exit of the eighth pair of nerves, being 
the most anterior of all, are situated on either side, in well-marked depres- 
sions or pits, some 7 millimetres apart. Back of these and nearer to- 
gether are the minute precondyloids, looking forwards and outwards for 
the passage of the hypoglossal nerves. These last foramina are just an- 
terior to the border of foramen magnum ; this latter aperture is of good 
size, comparatively, having an tero -posterior and transverse diameters 
of 3 millimetres each, with an additional millimetre for the oblique 
diameters, making the latter 4 millimetres each. It is snbcircular in 
outline, its anterior rim passing around a depression that lies jnst in 
front of the condyle, giving the latter the appearance of jutting out 
into the foraminal space. The condyle is nearly sessile, having the 
merest trace of a neck, hemispheroidal in form, with an horizontal 
and average diameter of .5 of a millimetre. Above and midway, later- 
ally, the borders of the foramen are encroached upon by the petrosal on 
either side, giving it rather a constricted appearance ; from these points, 
as we follow the posterior .moiety of the foraminal periphery, we find it 
to be grooved, each groove ending posteriorly within a millimetre of each 
other, in a minute foramen that traverses the internal table of the cra- 
nium upwards, outwards, and forwards for a short distance, thence to 
arch around, as a sinus, the epencephalic fossa, to meet in the longitu- 
dinal sinus coming from above. This arrangement obtains in the Cor- 
vidw, and some other families, where it is more strongly marked. The 
diapophyses of the occipital vertebra are in a plaoe but a little lower 
than the basi-sphenoid 5 they form, as is quite common, the horizontal 
floor of the cavity of the otocraue, and blend with the surrounding bones. 
A moderately well-marked "cerebellar prominence" occupies its usual 
site in the middle line; no openings or foramina are ever to be discovered 
either at its summit or laterally, as seen in some other birds (Anatid.ce, 
Strigidce). It divides the shallow temporal fossae that slope away from 
it on either side, and varies somewhat in size in different individuals. 
From the upper region of the ear and the superior boundaries of the 
temporal fossae to the line of that psuedo-articulation, the fronto-man- 
dibular, this bird's cranium is remarkably smooth, and of a clear white, 
and, owing to the extraordinary amount of diploic tissue, possessing a 
peculiar translucency. The median furrow is only well marked as it 
passes between the orbits ; the superior peripheries of these cavities, as 
constituting one of the boundaries of the surface under consideration, 
are sharp at first, rounding as they include the lacrymals, and entirely 
devoid of any notches or indentations. As is usual, all sutural traces 
are absent (PI. IY, Fig. 25). The transverse line of the fronto-mandib- 
ular juncture is slightly concave backwards along its middle third, the 
extremities sloping a little downwards and backwards. The joint mo- 
tion is only moderately free. No well-marked suture defines its exact 
locality, as in Harjwrhynchus and others. The bones that go to form 



630 GEOLOGICAL SURVEY OE THE TERRITORIES. 

the superior mandible, both above and below, are mutually confluent at 
all their usual points of contact and articulations, with complete oblit- 
eration of their original borders. The nearly perpendicular nasals on 
either side form the anterior boundary of a triangular opening, of which 
the lacrymals and maxillaries form, respectively, the posterior boundary 
and base. These triangles are not complete, insomuch as the lacrymals 
do not meet the infraorbital bars at the inferior and inner angles. They 
lead into the rhinal vacuity on either side. It must be borne well in 
mind by the reader that hi describing the upper mandible in the skulls 
of all birds, it invariably presupposes the removal of its horny in tegu- 
mental sheath that it wears during life, and gives to this portion of the 
cranium a vastly different shape. Either tomial edge is curved and 
quite sharp; their anterior mergence, or point of the beak, is decidedly 
rounded, and fully a millimetre in width. 

The superior mandible is rather broad at its base: the culmen. origi- 
nating in a flattened space just anterior to the fronto-mandibnlar artic- 
ulation, is rounded throughout its extent and gently curved downwards, 
while below, the line joining the middle points of the bases of the triangles 
above mentioned, averages 7 millimetres in length. The sides of the in- 
ter-maxillary are smooth, presenting only occasionally a row of very mi- 
nute foramina for examination ; sometimes a faint suture shows itself on 
either side, extending almost down to the nostril, between this bone aud 
each nasal. Beneath, the palatine fissure is broad and rounded ante- 
riorly, the roof of the mouth beyond being gently concave and grooved 
mesially for its entire length, and marked by a few foramina. The ex- 
ternal apertures of the nostrils are quite large, nearly elliptical in out- 
line, approaching each other within less than .5 of a millimetre above. 
Their borders, formed by the nasals behind, are sharp: anterior, more 
rounded. The major axes of these openings average 4 millimetres, 
the corresponding minor axes 3 millimetres. The planes passed through 
their peripheries look upward, outward, and forward. The nasals are 
fan-shaped, both above and below, the expansion being slightly twisted, 
in order to accommodate themselves to the form of the bill. The broad 
lacrynjals, assisted by the prefrontal, effectually separate the orbital 
vacuities from the rhinal chambers. The latter are more than usually 
open, owing to the size in the skull of the various apertures leading into 
them from without, already described, and devoid of all septa or bony 
offshoots, although the prefrontal, intermaxillary, and palatines to- 
gether occasionally develop irregularly formed ethmo-turbinals. that 
extend into this space from behind and afford the necessary surface for 
the pituitary membrane. But there is nothing that has the slightest 
semblance to an osseous septum narium. The anterior olfactory fora- 
mina, narrow slits one millimetre long, are found between the lacrymals 
and prefrontal, close to the vertical septum of the latter: their outer ex- 
tremities being the superior, they are seen to look downward and for- 
ward as they open into the nasal cavities from the bases of the concavi- 
ties formed by the bones above mentioned. 

The orbital cavities are capacious, having rather a forward look : at 
the same time they look a little downward. Their limiting borders are 
ovate in outline, with the greater end backward, being incomplete be- 
low. Anteriorily the septum that divides them is remarkably entire and 
of considerable thickness : posteriorly and above there exists quite a 
deficiency, of a shape shown in PI. IV. Fig. 22 : this is situated just in 
front of the large quadrilateral rhinencephalic foramen, and allows a good 
passage from either orbit into the brain-case. The same condition ob- 
tains below with the opening for the exit of the optic nerves, only the 



snuFELDT.J OSTEOLOGY OF THE EREMOPHILA. 631 

latter is much smaller and quite circular; to its outer side there are 
several miuute foramina that lead directly into the brain-case. The 
groove for the first pair is distinct anteriorly on either side, and opens 
into slits between the prefrontal and lacrymals, similar to those described 
when speaking of them in connection with the nasal cavities. These, 
the anterior rhinal foramina, together seem to be the hoinologue of the 
" cribriform plate' 7 of anthropotomy. The anterior wall of an orbit is 
formed by a lacrymal ; this bone is larger than usually found in avian 
crania of this size. It is quadrilateral in form, concave posteriorly, thor- 
oughly confluent with the frontal, nasal, and ethmoid, but not coming 
in contact either with the palatines or suborbital style. About the mid- 
dle of its outer border it presents a rounded notch for the lacrymal duct. 
Its anterior surface, forming the posterior wall for the rhinal vacuity, 
is undulating, though generally convex. 

The superior wall of the orbit is narrow, gently concave, and formed 
as usual by the frontal. It looks downwards and outwards and merges 
into the orbito-cranial septum behind, conformably with the shape of the 
cavity under consideration. The posterior wall of the orbit presents 
quite a number of interesting points for examination. Internally and 
above we find the posterior rhinal foramen, and below it the foramen 
opticus, already described. In addition to other minute openings men- 
tioned above, we have the foramen ovale, occupying a lower plane than 
any of the others, and situated more external to them, being almost di- 
rectly behind the orbital process of the tympanic. Above it we observe 
a thin circular convexity, indicating the locality of the mesencephalic 
fossa; this sometimes develops at its outer border a sharp, vertical, 
osseous spine or plate, that points downwards, forwards, and inwards 
into the orbital cavity. Still beyond this, outwardly, we find another 
X^rocess, or rather two processes combined, with an elliptical foramen 
between them, placed vertically. The inner portion consists of a square 
lamina of bone, looking upwards and forw r ards ; the other smaller and 
outer portion is a trihedral spine that descends, apparently from the 
frontal, to meet its external margin. The arrangement gives to the en- 
tire posterior wall a certain facing, directly forward, forcing upon the 
cranium of this little bird an aspect peculiar to another family, from 
which it is far removed — fche Strigidce. 

The osseous floor of the orbital cavity is always more or less imperfect 
throughout the class, and is here formed by the customary bones, the 
tympanic, pterygoid, slightly by the lacrymal, and limited externally 
by the malo-maxillary squamosal bar. 

The palatines nowhere come in contact wilh each other, and the pala- 
tine fissure is very wide, broadly rounded at both ends. The anterior 
extremity of each of these bones articulates in the usual manner, with 
the maxilliary and intermaxillary, the joints being immovable ones. 
Back from this point as far as the under surface of the lacrymals, on 
either side, they are but very slender, straight, and horizontally flat- 
tened little bones, without plate or process; at this latter point they 
suddenly expand into quadrate posterior ends, each slightly inclined 
downwards towards the median plane, throwing out a thin, nearly ver- 
tical plate for articulation with the fan-like and anterior ends of the 
pterygoids, while mesially they develop two other slender horizontal 
plates, the superior one being prolonged forward as a fine spicula of 
bone to meet the ethmo turbinal mass, as above described. 

They lightly touch the rostrum of the sphenoid, in company with the 
pterygoids, forming the usual arthrodial joint at this point in avian 
structure. Above they are smooth, look upwards and outwards, and 



632 GEOLOGICAL SURVEY OE THE TERRITORIES. 

form a portion of the floor of the orbit on either side. The union 
among tli- pre sphenoidal process and prefrontal plates is com- 

plete, all sutnral traces having disappeared, and the included bones form 
the interorbital septum as already described. The zygomatic style, 
very slender, straight, and throughout its continuity nearly of uniform 
calibre, descends from before backwards from its maxilliary articulation 
to the tympanic, about 4 millimetres, the skull being horizontal. 

The coalescence among its three original element is unusually per- 
fect. The anterior horizontal expansion is very slight, being crowded 
towards the intermaxillary osseous toniium on either side by the widely 
separated pala tines. Its posterior extremity is club-shaped and turned 
upwards, bearing on its inner aspect a hemispheroidal articular facette 
for the cotyloid cavity of the tympanic. 

In no single articulation found in the skeleton throughout the class 
does there seem to be more variation in plan, to meet the same end 
and carry out the same function, than we find in the ptery go-pala- 
tine with the rostrum of the basi-sphenoid. In our present subject, 
as in Pica and Corrus and many others, this extremity bears a thin 
expansion that articulates by its anterioi edge with the palatine 
plate and neatly grasps the rounded and inferior side of the ros- 
trum, the two bones not usually coming in contact. The shaft of 
the pterygoid also slightly expands horizontally just before this artic- 
ular surface is developed, more particularly in the angle between 
the two, adding greatly to the sta L_i;h of the bone, and somewhat to 
the floor of the cavity of the orbit. The angle of divergence of the 
pterygoids in the present instance is exactly 45°: the intertympanic 
chord. 7 millimetres. The shaft of this bone is comparatively slender, 
prism oidal in form, somewhat twisted, and develops among the older 
birds sharp projecting edges. The enlarged tympanic extremity bears 
a subelliptical articulating facette, that glides upon a similarly formed 
surface surmounting the ptyergoidal process at the base of the orbital 
process of the -ponding tyinpanie element. These two little bones 

are well separated from the basi-sphenoid, and never any evidence of 
the development of pterapophysial processes ie to be observed. As is 
generally, though by no means universally, the case among birds, the 
mastoid process of the tyr>._ in this Lark is distinctly bind, each 

limb presenting for examination at ite rz:iemityan elliptical convex 
facette for articulation in a cup-shaped cavity intended for its reception 
in the roof of the aural vacuity. Of the tw irfaces, the outer and at 
the same time the aDterior looks outwards, forwards, and upwards, while 
the inner and posterior one. surmounting the shorter limb or bifurcation, 
looks backwards and upwards. These two projections of the mastoid 
process are further separated posteriorly by a deep non-articular de- 
pression. The orbital process is ^vell developed, long and slender, ter- 
minating in a knobbed extremity, the whole extending well within the 
orbital space. It has at its base, internally, the facette for the ptery- 
goid already alluded to. This process is subcompressed from before 
backwards, and has throughout a gentle curvature upwards, having 
much the form of the thorn of the common rose, without its sharp point. 

There are two articular faeettes on the inferior side of the mandibular 
end, divided by rather a deep depression. Of the two, the inner is the 
larger and more symmetrical in form, being transversely ellipticaL The 
outer one seems to be borne on rather a constricted neck, having on its 
outer aspect the acetabulum for the hernispheroidal facette on the squa- 
mosal. The anterior surface of the body is smooth and triangular in_ 
outline ; the opposite and inner surface, somewhat similar in appearance, 



shufeuxt.] OSTEOLOGY OF THE EREMOPHILA. 633 

presents for examination, just below the mastoid process, a large, oval, 
pneumatic foramen ; other of these openings may exist in the depression 
on the posterior surface of the body of the bone already described. 

The inferior surface of the basi- sphenoid is convex outward, and slopes 
away gradually into the rostrum, anteriorly. The external orifices of 
the Eustachian tubes are extremely minute, as are the foramina for the 
entrance of the branches of the common carotid to the cranium. As 
already intimated when speaking of the pterygoids, there are no ptera- 
pophysial processes. 

The external aperture to the cavity of the otocrane is an elliptical 
slit, 1.5 millimetres wide at its widest part, looking almost directly for- 
wards, its lower end being the innermost or nearest the median plane. 
The mastoid, however, does not extend so far forwards but that in a 
direct lateral view we may see, through the opening, the funnel shaped 
internal orifice of the Eustachian tube. The stability of the ear cavity 
is here, as in many birds, highly enhanced by the presence of numerous 
osseous trabeculse, acting as struts and braces to its walls. 

An examination of the interior of the brain-case shows the fossae for 
the several cephalic lobes to be large — indicating a brain of good size 
for the bird. As already defined, the foramina for the first and second 
pairs of nerves are in each case single, and as a whole more or less oval. 
A constriction, however, takes place in their outlines at the middles, 
formed by the encroaching interorbital septum, so that, looking out of 
the cavity, the foramen in either case appears double, whereas a view 
from an orbit reveals the fact of there being but one opening in either 
case. The olfactory foramen is very large — in the dry cranium — the 
deficiency being made up by firm membrane in the living Lark. The 
minute openings for the carotids at the base of the pituitary depression 
are placed, as usual, side by side transversely. The posterior wall of 
the sella turcica is deeply notched. 

The longitudinal sinus is best seen along the superior and median 
crest, just before it arrives at the olfactory foramen. The middle fossa 
for the accommodation of the cerebellum is distinctly marked by long 
transverse concavities, admitting the rugae upon the lobe in question 
when the brain is in situ. With regard to the structure of this bird's 
cranium, we may say that it is largely cancellated, the intermaxillary 
and petrosal approaching nearest the compact variety of bone; this fact 
lends to this part of the skeleton a great lightness, and well-prepared 
skulls of this Lark are very pretty objects. 

The most remarkable feature to be observed, however, is the great 
amount of separation between the tables of the vault of the brain cavity, 
being fully a millimetre, and in some localities more, the interspace being 
filled in by quite an open diploic tissue. This condition we well know 
to be a striking feature in the anatomy of the Strigidce, but here is a 
bird that has the same arrangement as well marked, we believe, for its 
size, as any Owl in the North American fauna. The outline of the base 
of the cranium in Uremophila approaches the sector of a circle, a figure 
more or less true in all birds, and here, as in most others, the greatest 
departure from that figure beiag a too great convexity of the subtending 
arc. The length of the radius represented by the middle line is 3.2 centi- 
metres, the interrympanic chord, including the bones, being 1.4 centi- 
metres. We will only mention here, in regard to the free osseous ele- 
ments of the sense capsules, that the sclerotals retain their usual form 
and arrangement, numbering in each eye from thirteen to fifteen. The 
attachment among them is rather firm, remaining as shown in PL IV, 
Fig. 41, after a considerable amount of maceration. The ossicula auditus 



634 GEOLOGICAL SUEVEY OF THE TERRITORIES. 

are also present, but a lens of some power is required to study their 
form and arrangement. 

The hyoid arch — (PI. IY, Fig. 37, seen from below). — This, the hsernal 
arch of the parietal vertebra, in no way deviates in this little Lark from 
the usual ornithic characters possessed by it among living birds, in being 
freely suspended beneath the cranium and acted upon by certain mus- 
cles. The glosso- and cerato-hyals seem to be confluent, and the bone 
thus formed consists in two narrow little affairs, that for their anterior 
two- thirds run alongside of each other with a greater or less intimacy, 
to have their tips slightly diverge anteriorly. Posteriorly the ends have 
a still greater amount of divergence, and at the junction of the middle 
and posterior thirds there is a transverse bony bridge, that bears the 
facet te for articulation with the basi-hyal behind. Scarcely any antero- 
posterior curvature exists. The posterior tips overhang the articula- 
tion of the thyro-hyals with the confluent basi- and uro-hyal. As we 
have never examined the tongue of the young of Eremophila, we may 
be in error in saying that the glosso- and cerato-hyals are confluent, as 
the bones we have just described may be the cerato-hyals alone, the 
glosso-hyal being entirely in cartilage. The posterior tips of the cerato- 
hyals have an expansion to accommodate the articulation referred to, 
bearing on either side small, elliptical, articular surfaces, looking back- 
wards and outwards for the heads of the hypo-branchial elements of 
the thyro-hyals. 

The bone is subcompressed from above downwards, the uro-hyal being 
produced behind by cartilage, or rather tipped by that material, while 
the articulation at the anterior extremity of these confluent bones is 
hidden from view in the superior aspect of the arch by the glosso- and 
cerato-hyals ; and, as is common, the inferior lip that the basi-hyal lends 
to this joint is the longer, and protrudes forward. 

The hypo- and cerato-branchial elements of the thyro-hyals are very 
long, slender, up curved little bones, produced posteriorly, as is the uro- 
hyal, by cartilaginous tips. 

The shaftlets of these delicate- elements are slightly flattened from 
above downwards, as are their articular heads. The free extremities 
have a tendency to curve inwards a little, or towards the median plane, 
as well as upwards. 

The loicer mandible — (Plate IV, Figs. 22 and 29). — Uremophila is an- 
other example exhibiting the non-approximation of the tomial edges of 
the mandibles in the dry skull, this feature being more often absent 
among Grallatores and many of the Natatores, where these edges come 
in contact with almost an equal amount of exactness as where the bill 
is armed with its horny theca. 

The lower mandible of the Horned Lark seems to be, in point of struct- 
ure, composed almost entirely of compact tissue, and, owing in addition 
to the thorough coalescence of its primary elements, a very firm and 
strong bone. Sutural traces, the indicators of the boundaries of pris- 
tine segments, have entirely disappeared, and no one would ever suspect, 
in examining it, the presence of nine original parts, were he not familiar 
with avian osteology or had the opportunity of dissecting the young. 
The inferior surfaces of the articular ends are on a level with the major 
part of the under rim of the rami, but they are well below the coronoidal 
elevations on either side. They present superiorly the usual undulatory 
surface to meet and articulate with the condyles of the tympanies. 
Below appears a longitudinal ridge, due to the extension upon that side 
of the ramal edges. A knob like process projects behind, and the true 
articular processes are sharp and rather long. They are directed in- 



6HUFEI.DTJ OSTEOLOGY OF THE EREMOPHILA. 635 

wards, upwards, and then forwards, having the usual pneumatic foramen 
above and near their pointed extremities. The superior margin of the 
inferior maxilla starts at once from each articular surface, to rise by a 
moderate angle to the representative coronoids, a distance of 4 milli- 
metres ; it then falls gradually to the rounded and anterior termination 
of the bone. It exhibits about its middle, on each side, a long but very 
low convexity, the corresponding shallow concavities being between them 
and the coronoidal elevations. The u coronoids" are marked by deep 
groovelets with raised borders that extend forwards and downwards as 
far as the interangular vacuity. 

The inferior boundary of the bone, as already stated, rises on each 
side in the inferior articular surfaces, to ascend first for two-thirds of 
its extent on each ramus, then to fall at about an equal angle, to sweep 
round and form the anterior and curved termination in the dentary ele- 
ment. The median line on the dentary segment averages 5 centimetres, 
this portion of the bone being quite thick and concave above, convex 
below. The general surface, both inside and out, between the bounda- 
ries just defined, is in each case depressed, smooth, and translucent until 
we arrive at the solid dentary portion, where we find it marked by a 
row of minute pits. Of some dozen or more lower maxillae before me, 
one of the most striking differences existing among them seems to be 
the variation in size of the interangular vacuity or foramen. This is 
elliptical in outline with the major axis of the ellipse in the long axis of 
the bone, and in some specimens squarely meet the raised ramal borders 
within, while in other individuals, even though the bone be larger, this 
foramen is markedly smaller. A large concavo-convex sesamoid is found 
between the tympanic and articular end on each side. The long axes of 
these bones are placed vertically, and their concave surfaces look for- 
wards. They are attached to the middle of the pointed articular pro- 
cesses behind by a delicate ligament, and above by the same means ; by 
a somewhat broader attachment to the squamosals and tympanies, pos- 
teriorly. 

Spinal column, cervical portion — (PI. IV, Figs. 22 and 35). — In making 
a study of the vertebral column of this Lark, the student will find that 
he will be materially assisted if he make use of an engraver's eye-lens, 
or, better still, one of the low-power objectives of a good microscope, as 
some of the points for examination are rather minute, and are not so 
easily or satisfactorily demonstrated by the unarmed eye. The cervi- 
cal portion of the column is composed of thirteen vertebrae ; these enjoy, 
from theatlantal throughout the entire series, a perfectly free move- 
ment among each other by their several articular surfaces ; and some 
form of the sigmoidal curve, characteristic of the bird-neck, is invaria- 
bly preserved during life and action. We find, too, the majority of the 
salient points pertaining to these segments described by ornithotomists, 
present, and strongly marked, and the chief functions of this jointed and 
bony isthmus well carried out — as affording protection for the myelon 
in its passage from the brain to the body below, and the vessels from 
their centre to the brain above. The neural canal, beginning in the 
atlas as a transverse ellipse, rapidly becomes circular, retaining this 
form throughout the tube, only to resume the elliptical again in the last 
two or three segments, where in the thirteenth it seems to be of a lar- 
ger calibre than at the cranial extremity, the ellipse still. being placed 
transversely. 

The usual processes often of these vertebrae, the third to the twelfth, 
inclusive, afford protection to the vertebral artery and sympathetic 
nerve. By an apparent contraction of the parapophyses in the twelfth, 



636 GEOLOGICAL SURVEY OE THE TERRITORIES. 

the canal is open laterally in this segment. It is confined to the anterior 
third on each side of the vertebrae enumerated, and is exceedingly small 
throughout its extent: its largest calibre "being at its commencement, 
its finest in the tenth or eleventh. Among the long vertebra? in the 
middle of the neck the anterior entrances of the vertebral canal are 
ellipses placed vertically. They become more circular as we approach 
the thoracic end of the chain. On the eighth vertebra, mesialiy. and 
beneath anteriorly, we find, bounded on either side by the parapophysial 
processes, the commencement of the interhyapophysial groove or canal 
for the carotid artery. It extends through the fourth vertebra with 
about an equal amount of distinctness and depth.. 

A neural spine is feebly developed upon the axis posteriorly, this 
process becoming more strongly marked on the summits of the next 
three succeeding vertebrae, the remainder of the cervical segments being 
devoid of this feature, though we have occasionally found an evident 
attempt at its reproduction in the ultimate cervical. The nethermost 
portion of the pseudo-centrum of the first vertebra has been considered 
to be the atlantal hypapophysis. Be this as it may. the hypapophysis 
of the axis certainly has a much greater claim to be termed a process, 
while on the third and fourth segments this spine constitutes one of 
the most marked features of the vertebra, being a longitudinal and quad- 
rate lamina of bone, equally well developed on the two vertebrae in ques- 
tion, directed immediately forward. In the case of the fifth cervical the 
hypapophysis has again degenerated to a minute median point, to be 
entirely obliterated in the sixth. At the ninth it again makes its ap- 
pearance as a delicate and flattened plate at the anterior margin of 
the vertebra beneath, at the point at which in the carotid canal it is first 
seen in the eighth. In the remaining ones it is prominently developed 
and directed forwards from the median plane in each vertebra as a quad- 
rate lamina. It is usually triplicate in the last, but does not arise from 
a common stem, as in other birds. 

Parapophysial processes appear as lateral spines first on the third 
cervical: in the middle of the series they are very long and delicate, 
being parallel with the centrum of the vertebra to which they belong. 
They become markedly suppressed near the termination of this division 
of the spinal column. 

Anterior and posterior zygapophyses retain throughout the cervical 
vertebrae their most common ornithic features : in the middle of the 
neck the postzygapophysial processes are long and bent slightly towards 
the neural canal, leaving quite an extensive lozenge-shaped space be- 
tween them in this region where the chord is unprotected by bone: the 
interarticular facets among the centra likewise retain their most com- 
mon avian characteristics. The bodies for the most part seem to be 
slightly compressed from side to side, with a faint inferior median crest. 
The fourth vertebra has a delicate and outwardly-arched interzygapo- 
physial bar. that includes within it an elliptical foramen on each side 
of some size. This bony connection in the third vertebra nearly fills in 
the interzygapophysial space, a very minute vacuity alone remaiuing. 

All the cervical vertebrae appear to be pneumatic, but the foramina 
in some of them are excessively small and difficult of detection. 

The bony cup of the atlas is not usually pierced by the odontoid 
process of the axis, but one cannot but wonder, this cup being less 
than half a millimetre across, that the skull, so very large as compared 
with its tiny occipital condyle, should not be subject to frequent dis- 
locations: this undoubtedly would be the case were not the occipito- 



shufeldt.j OSTEOLOGY OF THE EREMOPHILA. 637 

atloid articulation so thoroughly re enforced by the thick muscles that 
surround it. 

A square bony plate projects from below in the atlas, more anterior 
than any other part of the bone, that covers the atlo-axoid articulation 
in front. 

The arch that connects the neurapophyses is broad and smooth, and 
assists greatly in the protection of the myelon between the two bones. 

The odontoid process on the axis is concave in front, flat behind, with 
a roundish summit. It averages one millimetre in length, and is directed 
slightly backwards. The articular surface at its base is reniform in 
outline, the centrum that supports it being contracted below. The 
postzygapophyses show faint traces of anapophysial tubercles; these 
are better marked in the latter cervicals. The last or thirteenth verte- 
bra has freely suspended from beneath each diapophysial articular sur- 
face a rudimentary pleurapophysis that averages about two millimetres 
in length. These little bones represent the only true cervical ribs, though 
we must admit here that in several individuals we found the first pair 
of dorsal pleurapophyses unconnected with the sternum by the usual 
haem apophyses, and ending in pointed extremities. Should such a speci- 
men alone be examined, we would have to recognize fourteen cervical 
vertebrae, the last two bearing free pleurapophyses, but the common rule 
must dictate here as elsewhere, and the condition just mentioned be 
reckoned as the exception. 

Dorsal vertebra?, vertebral and sternal ribs, sternum — (PI. IV, Figs. 22, 
24, 27, and 38). — The number of vertebrae devoted to the dorsal portion 
of the spinal column in Mremophila seems to be invariably Jive. They 
are easily detached one from another, and after ordinary maceration of 
the skeleton drop apart almost as readily as the cervical vertebrae, so 
that during life there is at least quite a little amount of free movement 
among these bones. 

The neural canal, as it passes through this series, starts with the 
transverse ellipse as we left it in the last cervical, in the vicinity of the 
dorsal expansion of the myelon, to terminate nearly circular, and much 
diminished in calibre, in the ultimate segment of the sacral extremity. 

The neural spines form by their interlocking a continuous ridge 
above. The thickened crest of this ridge is produced by what we will 
call the arrow-head joint, a true schindylesial articulation to be found in 
many of the class. The superior margin of each spine becomes pointed 
anteriorly, extends forward, and is received into a fissure of the poste- 
riorly produced superior margin of the neural spine of the vertebra next 
beyond it. This arrangement has the appearance of so many little arrow- 
heads placed in simdar juxtaposition, and constitutes one of the elements 
of stability of the dorsal vertebrae in this bird. The open spaces remain- 
ing among the bodies of the spines below, between their produced crests 
and the several neural arches, are filled up by connecting ligament and 
membrane. 

The di apophyses of the dorsals are a very much horizontally flattened 
series. They are all slightly tilted upwards, the anterior ones being the 
broadest and shortest, and the ultimate one, by a gradual departure in 
this regard from the first, the narrowest and longest. In the middle of 
the series, moderately well developed and antero-posteriorly produced 
metapophysial ridges are found limiting the diapophyses externally; 
they do not reach from one vertebra to another. The pneumatic for- 
amina at the bases of these processes are very minute and scarcely dis- 
cernible by the naked eye. 

The inferior diapophysial facettes for the plcurapophysial tubercula 



638 GEOLOGICAL SCBVEY OF THE TERRITORIES. 

are c cue a ve- elliptical surfaces, with their major axes parallel to the 
median Hue. The anterior Dues are the more circular. 

The zygapophysial processes, to assist in the intimate proximity of 
these vertebra, are short and thick. The anterior ones look upwards 
and inwards, the reverse being the case with the posterior series, which 
latter develop pointed spines that overlap above, each in its tnrn. on 
either side, the vertebra next behind, at the base of the common neural 
spine. The longest of these processes are found anteriorly: they gradu- 
ally disappear as we near the sacrum. 

The first dorsal hypapophysis consists of three plates, arising from 
the centrum of the vertebra separately, and arranged as shown in Fig". 
38. On the second dorsal we find only a single quadrate plate in the 
median plane, directed forwards. It occupies a position at the anterior 
margin of the vertebra, bat is produced posteriorly as a low. thin lamina 
of bone, along the remainder of the centrum mesial to the raised and 
posterior margin. The third vertebra takes it up in this form, and it 
is thus passed along the series, constituting a continuous hypapophysial 
ridge, intersected by the expanded anterior and posterior borders of the 
centra. 

The articular surfaces among the bodies retain their usual characters. 
They extend into the ridge just described. The centra of the dorsal 
vertebra are somewhat compressed in a slightly increasing degree from 
before backwards: each lateral and anterior margin supplies a nearly 
circular parapophysis for the pleurapophysial cafitula, while at points 
on the posterior margins in the same plane we n d the major share of 
the notch, which in coaptation of the segments constitutes the subcircu- 
lar foramina for the exit of the dorsal nerves. 

There is a free pleur apophysis for each dorsal vertebra, but the first is 
not always connected with the sternum by a sternal rib. as already de- 
fined: it sometimes has all the characteristics of a movable cervical rib: 
again, when it connects with the sternum, its tnemapophysis articulates 
rather high on the costal border (PL IV. Fig. 22). It may or may not 
bear an epipleural appendage. 

The vertebral ribs of this Lark articulate, as usual, by tubercula and 
capitula. with the dorsals, meeting par- and di-apophyses in the ordinary 
manner. The necks of the ribs in the middle of the series are the long- 
est, and often we find among the ultimate ones a slight projection be- 
yond the tubercle, that is received in a corresponding notch at the outer 
border of the diapophysis it meets. There is but little difference in the 
width of these fiat bones; perhaps the anterior ones have rather the 
advantage in this respect. Minute apertures, to allow the air to enter 
their bodies, are observed in the usual localities. 

The laterally viewed curve of a dorsal rib is barely sigmoidal: viewed 
from in front it approaches a portion of the curve of an arc of an ellipse. 

A ridge continuous with the neck is carried down the inner aspect of 
each bone, to gTadually disappear near its middle. The lower extremi- 
ties of these ribs are slightly enlarged, to afford space for articulation 
with the sternal ribs: the surface is convex. 

The epipleural appendages of the dorsal pleurapophyses are conflu- 
ent with the posterior edges of the bones, and situated below their mid- 
dles. Occasionally the one in the middle of the series has sufficient 
length to overlap two ribs: in young birds of this species they are much 
shorter, and the best-developed ones show an angle on their inferior 
borders j nst after leaving the rib. as if they had left that bone with the 
original intention of proceeding downwards and backwards at a gentle 



shufeldt.j OSTEOLOGY OF THE EREMOPHILA. 639 

angle, but suddenly changing mind, proceeded directly upwards and 
backwards at an equal angle; hence the condition alluded to. 

When the first dorsal rib articulates with the haemal spine below by 
the intervention of a sternal rib, this latter bone is quite small and del- 
icate, averaging about 3 millimetres in length, and but slightly curved. 
The remaining dorsal haemapophyses become longer and more curved 
as we follow them backwards. They are all flattened from side to side, 
their lower extremities being abruptly twisted at right angles with their 
shafts, enlarged, and terminating in a flattened articular surface for the 
costal border of the sternum. These articular surfaces are dumb-bell 
shaped, i. e., contracted in their middles. The upper ends of these sternal 
ribs are also enlarged and laterally flattened for articulation with the 
vertebral ribs. These latter enlarged ends are sometimes larger, some- 
times smaller, than the extremity of the pleurapophysis they meet. 

If we accept as true the old vertebral theory of Oken, Goethe, and 
Owen, a theory that holds that the skull of all vertebrates is composed 
of three, four, or more modified vertebras, that these vertebrae in evolut- 
ing from the very earliest forms in which a bony segmented column 
appeared, that during this evolution many metamorphoses took place 
in the skull, such as the restriction of the notochord to the first or occi- 
pital vertebra, the mode of development of the elements themselves, 
whether by cartilage or membrane, and the appropriation by these 
" cranial vertebrae" of bones that did not originally belong to them, that 
we cannot positively say what may become of the cranial extremity of 
the spinal column of Amphioxus, or some of the haemal arches in the hag 
and others, then we must recognize in the sternum of birds, developed 
as it may be, the confluent haemal spines of the dorsal series of ribs ; 
and in it, in its maturity, see one of the most interesting bones to con- 
template, it being one of the most diversified in form in the bird skeleton. 
Owen styled the type of this bone, as found in the Lark now under 
consideration, "cantorial" (Anat. and Phys. of Vert., Vol. II, p. 20). 
It is certainly typical of the suborder Oscines, as far as American orni- 
thology is concerned ; good examples as testifying to this I have now 
before me, in the haemal spines of Turdus migratorius, Ampelis garrulus, 
Mimns polyglottus, Lanius, and many others. 

In Eremophila the sternum is very light and delicate in structure; so 
thin is it in some individuals that we find deficiencies occurring, usually 
in the body, as foramina of no mean size (1.8 millimetres). Its outer 
surface, indeed the entire surface of the bone, has the appearance as if 
it were venated, the solid bony veins being thicker and more opaque 
than the general surface of the bone, and branching from the various 
borders. 

The carina is moderately well developed, measuring in the vertical line 
below the coracoidal groove 9 millimetres. Its inferior border, expanded 
behind, is rounded and somewhat thickened; this thickening disappears 
on the anterior border, which is sharper and continuous with a conspicu- 
ous crest on the front of the manubrium. 

The carinal angle, with an aperture of 70°, is quite prominent and 
produced anteriorly. Just within the anterior margin of the keel we 
find a rather prominent carinal ridge, its lower extremity branching 
backwards, and by its ramifications taking part in the superficial vena- 
tion referred to above. The keel arises abruptly from the inferior and 
median angles formed by the sides of the body where they meet mesiad. 

The xiphoidal prolongation is profoundly notched once on each side. 
These notches have the outlines of isoceles triangles, with their angles 
rounded, and apices but a short distance from the costal borders. These 



640 GEOLOGICAL SURVEY OF THE TEEEITORIES. 

deep indentations of the xiphoid give rise on either lateral sternal bor- 
der to a long, stout process, extending backwards and outwards, with 
dilated extremity. 

The outer surface of the body of the sternum presents for examination 
well-marked pectoral ridges, and, running from the bases of the xiphoidal 
processes to tbe outer angles of the coracoidal depression, clearly defined 
subcostal ridges. 

The manubrium is a prominent, superiorly bifurcated, trihedral pro- 
cess, jutting out from a substantial base iu the median plane, forwards 
and upwards, from the angle formed by the coracoid groove and the 
front border of the carina. At its base internally there is an extensive 
oval pneumatic foramen. Its bifurcations are rounded, and give attach- 
ment at their extremities to firm ligaments, that pass directly to the 
coracoidal capitula above. 

The groove for the coracoids is strongly marked and continuous in front, 
exteudiDg from costal process to costal process; its boundaries form the 
thickest and stoutest part of the bone we are describing. 

The costal processes, possessed of broad bases, arise as thin but prom- 
inent lamina, upwards, forwards, and outwards, terminated by flattened 
summits. Their posterior margins bear the costal facets for the articu- 
lating ends of the sternal ha?inapophyses. 

The sides of the body of the sternum on its ventral aspect make an 
obtuse angle with each other. The line of meeting in the mesial plane 
is quite evident : its anterior half is the seat of a row of various-sized 
pneumatic foramina. There are upon each costal border five, sometimes 
six. transverse facettes for the sternal ribs : the shallow depressions 
ainoDg them are scantily supplied with pneumatic foramina. 

The mid-xrphoidal border, in which the keel terminates posteriorly, 
is thickened; its other boundaries are sharp, with raised ridges below, 
just within their edges. The greatest length of the sternal body is a 
little more than two centimetres, and its greatest width a little more 
than one centimetre, the last measurement taken to the rear of the cos- 
tal processes. 

Sacral vertebra and ribs, pelvis, coccygeal vertebrae — (PI. IY, Figs. 22, 
23. and 28). — The first sacral vertebra has become thoroughly confluent 
with the ossa innominata on either side and with the vertebra behind 
it. Its diapophyses seem to have spread out upon theunder surface of 
the ilia, combining with them, for we observe that the first sacral pleu- 
rapophyses articulate in the ordinary manner with the transverse pro- 
cesses and the parapophyses, the tubercula being situated just near the 
outer iliac borders. This rib may become, as a rare event, confluent 
with the pelvis, but is usually free. Its haemapophysis is the longest of 
the series, and the articular facette on its lower extremity meets the 
last facette upon the sternal costal border. This sacral pleurapophysis 
may possess an epipleural appendage, though it seems to be the excep- 
tion. 

The second sacral rib is a delicate hair-like bone of uniform thickness, 
that does not show any decided tubercle, merely, after leaving the ver- 
tebra, coming in contact with the under surface of the ilium, on each 
side, for the entire interspace between the tubercle and head. It, too, 
may become confluent with the pelvis on its lower surface. 

Extending downwards and forwards by a gentle curve, it meets its 
haemapophysis through a miniature articulation. This latter style ar- 
ticulates along the posterior border of the sternal rib of the first sacral 
pleurapophysis, never reaching the costal border, and the second sacral 
nb never bears an epipleural spine on its posterior border. 



shufeldt.] OSTEOLOGY OF THE EREMOPHILA. 641 

The sacral vertebrae are invariably confluent throughout the chain in 
the pelvis of the Horned Lark; indeed, it is only by a process of staining 
this compound bone, and the aid of a strong light, that they can with 
any satisfaction be counted. There are eleven of them ; exceptionall} , 
twelve. 

The neural canal, circular at the outstart, shows the usual pelvic swell, 
chiefly anterior to the acetabula, conformable with the ventricular dila- 
tation of the myelon in that locality. The exit of this tube distally is 
likewise nearly circular. The foramina along the bodies of the centra, 
in the vicinity of the dilatation referred to, are double and placed one 
above another, for the separate egress of the roots of the pelvic plexus. 

The anterior aspect of the first sacral vertebra presents every element 
and process requisite for articulation with the ultimate dorsal segment. 
It is largely overshadowed by the ossa innominata. Opposite the iliac 
contraction, in the neighborhood of the fourth and fifth sacrals, these 
vertebrae throw out their par- and di-apophysial processes far enough to 
meet and brace the iliac bones. We do not meet with such braces again 
until arriving opposite the acetabula and beyond, where the parapo- 
physes project upwards and unite with the ouier margins of the trans- 
verse processes, the ilia articulating with the free and united borders. 

Foraminal deficiencies not unusually occur among these processes, 
more particularly between ttie last two sacrals, where they seem to be 
constant, though of varying size and shape in different individuals. 

The last sacral vertebra is compressed from above downwards, retain- 
ing, however, all the elements required in articulation with the first and 
much-modified coccygeal vertebra. 

Viewing the confluent sacral vertebrae, or the "sacrum", from above, 
we find the united neural spines, as a vertical lamina, dividing the ante- 
rior interiliac space into two capacious ilio-neural grooves at that moiety 
of the bone. 

This common neural spine and the ilio neural grooves proceed back- 
wards until the gluteal ridge of the ilium curves outward to the antitro- 
chanter on either side. At this point the spine disappears with the 
grooves, the sacrum becomes nearly flat and spreads out, to gradually 
contract again before its ultimate dilatation in the diapophyses of the 
last vertebra. 

It is a well-known fact to the ornithotomist that the pelves of birds 
differ with respect to the ilio-neural grooves in one of two ways. As 
seen in this bird they are grooves, as the ilia do not meet the united 
spines of the vertebrae; they are very frequently, however, converted 
into canals in other orders, by meeting of the interested bones above. 
The condition as defined though, in the previous paragraph, as relating 
to Eremophila, seems to be characteristic of American Oscines. The sa- 
crum is slightly convex from before backwards on its upper surface, 
moderately concave along the confluent centra below. 

The pelvis of this bird is uncommonly wide and short, and the ischi- 
adic and pubic posterior extremities remarkably flared outwards. The 
anterior and inner angle of each ilium, apparently assisted by the dia- 
pophysis of the first sacral vertebra from beneath, is pointed ; the ante- 
rior border slopes backwards gradually, for a distance of 3 or 4 milli- 
metres, to the rounded anterior external angle of the ilium. Between 
this point and the acetabulum the iliac border is strongly concave in- 
wards, as is the surface of the bone above it, the preacetabular being 
included between this border and a well-defined gluteal ridge. The 
superior postacetabular iliac surface is nearly square in outline, convex, 
and equal to a little more than one-third of the bone. It is thin and 
41 H 



642 GEOLOGICAL SURVEY OF THE TERRITORIES. 

translucent, its outer and posterior borders receiving the greater share 
of osseous reenforcement, particularly in the vicinity of the antitro,* 
chanter. 

Posteriorly, the ilium, slightly aided by the ischium, is carried out 
from an ilio-iscbiadic, overhanging crest, as bony processes, with their 
points turned slightly inwards. 

These processes are strongly marked in another of our Oscines, Harpo- 
rhynchus rufus, a bird that has a strikingly angular and rather unique 
pelvis. 

The antitrochanter is subelliptical in outline, and faces downwards, 
forwards, and outwards. The articular surface is produced downwards 
as far as the cotyloid cavity, upwards slightly above the general sur- 
face of the ilium, and is bounded posteriorly by the ischiadic notch. 

The foramen at the base of the hemispherical cotyloid cavity has so 
far absorbed the bone that really scarcely anything remains of it beyond 
a cylindraceous acetabular vacuity, the internal and external apertures 
being circles of equal diameter, and the femur consequently relying 
almost exclusively upon its fleshy and ligamentous attachments to re- 
tain its head in the ring. 

Sutural traces of the margins of the pelvic bones as the components 
of this osseous ring have entirely disappeared, having been obliterated 
during the pelvic consolidation. 

The ischium, for its major part, is like the ilium — very thin, more par- 
ticularly so at its free posterior borders ; joining with the ilium behind, 
it shuts off a large and elliptical ischiadic foramen, the superior arc of 
which is situated just beneath the ilio-ischiadic crest described above. 
The major axis of this ellipse is directed downwards and backwards. 

The posterior extremity of the ischium has an odd-appearing, foot-like 
termination, that is bent down to meet the pubis. 

This latter bone is an extremely slender style, that, immediately after 
assisting in the formation of the cotyloid ring, closes in a small, in fact 
the smallest of the group, subcircular obturator foramen behind; then 
running xiarallel with the ischium, by touching its further end, inclose, 
another long spindle-shaped vacuity ; it is finally produced beyond that 
bone by a pointed extremity, that curves backwards and inwards. 

It only remains now to say of the pelvis, as far as its internal aspect 
is concerned — after what we have said in regard to its extreme lightness, 
its translucency, its sacrum, and its borders — that, in general, superior 
convexities cause or create internal concavities, and vice versa. It is ca- 
pacious, and the various bones that compose it thoroughly anchylosed 
together. 

There are seven coccygeal or caudal vertebrce, rarely only six, and the 
pygostyle; they are in the skeleton so arranged and articulated that they 
have, as a whole, a gentle curve upwards, terminated by the quadrate 
" coccygeal vomer." 

These segments are all free, being easily individualized, even before 
maceration, by simple section of the ligaments that bind them together. • 

The subcircular neural canal, that passes through them, almost capil- 
lary in its dimensions, terminates without passing into the pygostyle. 

There is no hsemal canal developed, and indeed hypapophyses are 
found as stunted tubercles only on the last two or three vertebrae. 

A neural spine is developed on each, as a prominent and curved pro- 
cess, pointing forwards; this spine is wanting, however, on the last 
caudal. 

Of the lateral apophyses the transverse processes seem to be the only 
ones entitled to any consideration; these, as broad, flattened lamina, 



suufeldt.] OSTEOLOGY OF THE EREMOPHILA. 643 

extend from each vertebra, downwards and outwards, decreasing in 
width from before backwards; in fact, each vertebra in the coccygeal 
series becomes more and more rudimentary as we proceed in that di- 
rection. 

The articular facettes upon the centra start reniform, to terminate 
almost circular in the last vertebra; and the zygapophysial processes 
are exceedingly elementary in character. 

The pygostyle is parallelogramic in outline, articulating with the ulti- 
mate coccygeal vertebra by an imperforated cup-shaped depression, at 
the middle of its long anterior side. The edge of the bone above this 
point rests on the posterior border of the neural spine of the last caudal ; 
below it is free. 

The superior angle is more or less produced, and the posterior corner 
of the parallelogram is expanded laterally; this expansion is highly 
developed in many birds, as in Colaptes mexicanus and other members 
of the family Picidce. The caudal vertebrae are non-pneumatic in our 
present subject, whereas in the pelvis we find these foramina in their 
usual localities. 

The scapular arciv— (PI. IV, Figs. 22, 30, 32, 33, and 34).— This arch 
is very strong and perfect in this bird, as it is among the Oscines gen- 
erally. 

The bones can be easily separated from each other by maceration, 
though during life they are remarkably well strap ped together and to 
the sternum by their numerous ligaments. 

The scapula lies along the dorsum in its usual position over the dor- 
sal pleurapophyses, parallel with the vertebrae, with its posterior point 
touching the fifth one in the vast majority of the/specimens. 

Certain banes in all skeletons force upon us their resemblance to fa- 
miliar objects, and we know many of them have received their distinct- 
ive appellations through such likeness ; more particularly is this the 
case in the skeleton of man, where the bone we are describing is fre- 
quently termed the shoulder-blade, but how much more blade-like is the 
scapula in this Lark and many other birds, as far as shape is concerned. 
It is truly a miniature bony cimeter in EremopMla. This is not true for 
scapulae of all birds, however, for no one would ever be struck by such 
a resemblance while regarding the J-shaped scapula of Colaptes mexi- 
canus, or the straight, almost square-cut bone in some of our natatorial 
birds. 

In the Horned Lark the scapula is pointed and obliquely jtruncate be- 
hind for more than a third of its slightly dilated posterior portion, on 
the side towards the vertebrae. 

The outer border is reenforced by a rounded ridge for nearly its en- 
tire length, while the inner is quite sharp. 

The blade becomes stouter and subcompressed as we near the gle- 
noidal process; this broad tuberosity extends downwards, forwards, and 
outwards, and is crowned on its entire summit by a curved, subcircular, 
articular facet, that supplies rather more than one-third of the glenoid 
cavity for the head of the os humeri. 

The acromial process is bifurcated, and the clavicular head rests in the 
fork. The larger bifurcation is the lower, and both rest against the 
coracoid, on the inside and just below the head, creating the usual 
scapulo-coracoid foramen, which in this case is not very extensive. 

The scapula is pneumatic, and the foramina are to be found at the ex- 
tremity of the larger bifurcation of the acromial process, and in the 
notch between the two. 

The coracoid can boast of a very fair subcylindrical shaft between its 



644 GEOLOGICAL SURVEY OF THE TERRITORIES. 

head and inferior expansion. This flared extremity is quite thin out- 
wardly, stouter within, where it appears to be more of an extension and 
spreading of the shaft in its course downwards. Below there is a nar- 
row crescentic facet for the sternum, and at the upper edge of the ex- 
terior and thin side of the dilated end we find a notch, sometimes a fora- 
men, that appears to be constant. 

The upper extremity of the coracoid is an irregular tuberosity, con- 
sisting of a lower, inner, and smaller process for articulation with the 
clavicle, and an upper, superiorly convex head, that curls over mesiad 
to create a fossa, at the base of which we discover a group of various- 
sized pneumatic foramina. Anteriorly the head shows rather a well- 
marked process, into which the ligament coming from the born of the 
sternal manubrium, of the same side, is inserted. 

To the outer aspect, and below the head, is the reniform and vertical 
facet that, with the scapula and os humero-scapulare, goes to complete 
the glenoid cavity. 

The os humero-scapulare is a free bone, rather larger than the patella, 
found at the upper and posterior angle of the glenoidal process of the 
scapula. It is an elliptical disc, with a peg-like process extending from 
it from behind. The outer surface is concave and articular for the com- 
pletion of the glenoid cavity. This ossicle is held in position by various 
fibrous ligaments stretching from its borders to the scapular arch and 
the humerus. 1 

The clavicles are thoroughly fused together, forming one deeply U- 
shaped bone; their cylindrical and curved lengths support at the unioD, 
mesially and below, a long lamina of bone, in the median plane, that is 
directed upwards and backwards, parallel to the anterior carinal crest, 
to which it is united by ligament in the living bird. Their upper ends 
are expanded and placed in the skeleton flat-wise against the acromial 
process of the scapula and the head and the lower or clavicular process 
of the coracoid. The acromial process, through its bifurcation, partially 
grasps the hind border of this expanded end of the furculum, on either 
side. 

This bone seems to be non-pneuniatic, while the coracoids are hollow 
almost throughout their entire extent, having iu their composition very 
little cancellous tissue and a thin though firm, compact layer. The scap- 
ulae are hollow for some little distance into their blades, to be terminated 
by a cancellous structure, with an external and attenuated outer com- 
pact coat. . 

With the scapula arch in situ, we observe that the coracoids do not 
meet below in the coracoidal groove of the sternum, but approach only, 
on each side, as far as the periphery of the pneumatic foramen at the 
base and behind the manubrium. 

They are directed upwards, forwards, and outwards, at an angle of 
about 45° with the horizontal plane, the skeleton being erect; and, as a 
consequence, we find their upper ends further apart than any other part 
of the bone. 

The aperture between scapula and coracoid is in nearly a right angle, 
and the straight part of the inner scapular borders are parallel, their 
obliquely cut ends alone slightly turning outwards. 

A scapula is 2.5 centimetres long, a coracoid 3 centimetres, the inter- 
clavicular space above being 1 centimetre. 

In Turdus migrator ius we find the scapula shorter in proportion when 

^his little bone should be considered as a sesaraoidal auxiliary to the shoulder 
girdle, as it increases the articular surface of the glenoid cavity, and never be enu- 
merated as one of the bones of the pectoral limb. 



shufeldt.] OSTEOLOGY OF THE EREMOPHILA. 645 

compared with the other bones of the arch; the coracoids more de- 
pressed, i. e.j more in line with the sternum ; and the furculum in its 
direction backwards showing a gentler curve. 

The upper extremity — (PI. IV, Figs. 22, 31, 36, and 43). — The pectoral 
limb in Eremophila maintains the usual ornithic characters of a great 
number of the class, both in arrangement and number of the bones com- 
prising it. The skeleton arm has ten distinct segments; of these, we 
find one devoted to the brachium, two to the antibrachium, two to the 
carpus, one to the metacarpus, and four to the phalangeal portion of 
the manus. 

The bone humerus in this bird is more remarkable for its lack of curv- 
ature than anything else, being short and straight, as in others of the 
suborder among which our subject is classed and belongs. 

The head of the bone is broad and moderately flexed anconad, develop- 
ing only a very narrow and thin radial crest, which is bent for its entire 
extent toward the palmar aspect. This crest, answering to the " greater 
tuberosity" of anthropotomy, and giving attachment to the usual mus- 
cles, extends along on a straight line on the upper aspect of the shaft 
longitudinally for only about half a centimetre. 

The ulnar tubercle, or lesser tuberosity, makes up the thickened and 
proximal border of the confines of the pneumatic foramen ; a deep little 
pit on its palmar side or margin lodges the extremity of a strong liga- 
ment coming from the head of the coracoid of the same side, and mate- 
rially assists in keeping the head of the humerus in its socket. 

The elliptical and convex articulating facet of the head curls over an- 
conad, and from its middle a line runs down the bone for a short dis- 
tance, being one of the angular boundaries of this the trihedral extrem- 
ity of the bone. Quite a notch exists between the facet just described 
and the wall of the pneumatic foramen. This latter is on the under side 
of the head of the bone, snrrounded on its upper, proximal, and lower 
aspects by a firm bony wall, the lower and proximal parts of which are 
continuous with the smooth and otherwise unbroken surface of the ex- 
panded and palmar side of the head. 

The pneumatic fossa thus formed is deep, having at its bottom the 
foramen alluded to. Quite often the aperture is multiple, and vast 
differences in size exist, being very large in some individuals, nearly 
consuming the base of the fossa where it is found. From the lower 
boundary of the pneumatic enclosure another longitudinal line is seen 
on the proximal end of the shaft, limiting the anconal face of the trihe- 
dral end of the humerus in this direction. The palmar aspect of the 
head, broad and smooth, arches gradually inwards and towards the 
articular facet; it is also slightly convex from above downwards, sup- 
posing the bone to be in situ and in its position of rest, as we do during 
the course of our description. 

The shaft of the humerus is subcompressed from within outwards, 
smooth, and, viewing it laterally, it is barely convex above, by virtue of 
the ends being bent slightly down ; viewing it from above, we may say 
that it is almost straight. 

It retains its form until close under the expanded distal extremity, 
which is curved palmad. On the radial side of this end of the bone we 
find the transverse and convex elliptical trochlea below, for the sigmoid 
articulating depression of the ulna. This has inferiorly the quadrilat- 
eral internal condyle. 

The ulnar convexity is separated from the oblique tubercle for the ra- 
dius most effectually by a deep, well-marked, though narrow, notch. 

The oblique tubercle maintains its usual position as found on this bone 
in birds generally. 



616 GEOLOGICAL SURVEY OF THE TERRITORIES. 

The trochlea surface does not extend inwards very far : i. e.. does not 
pass over the end of the bone. Above it, and towards the proximal end,, 
appears a distinct and prominent external condyle. 

Anconad. this extremity of the humerus presents for examination the 
upturned internal condyle and a longitudinal tendinal groove, situated 
opposite the radial convexity, with intervening indentations. This ar- 
rangement lends to this aspect of the bone rather an uneven and tuber- 
ous look. The nutrient foramen, almost too minute to be observed by 
the naked eye, is found at the middle and inner aspect of the shaft. 

The radius is a long, delicate bone, with a bent and compressed shaft. 
A moderately well-expanded and circular head presents the usual eon- 
cavity for the oblique tubercle of the humeral trochlea, while below is a 
feebly marked ulnar facet and bicipital tuberosity; beyond this, again, 
the shaft develops a sharp, protruding edge, that extends nearly to mid- 
shaft and into the interosseous space. 

The distal extremity of this bone is spread transversely and curved 
downwards. It articulates with the upper surface and distal end of the 
ulna, and is lined above by very minute tendinal grooves. The outer 
border of this extremity presents a transverse lamina of bone that seems 
to be superadded to the dilated end. 

In articulation, the radius at first curves away from its companion, 
the ulna, to approach it again towards the carpal end. for about the outer 
third of the shaft, to remain with it until both arrive at the wrist. The 
distal border of the radius is transversely convex for an articular facette 
on the scapho-lunar. The idna is the main bony support of the forearm, 
and. indeed, its shaft is nearly equal in size and strength to that of the 
humerus itself, having the appearance of being the true continuation of 
the pectoral limb, so diminutive and slender is the accompanying radius, 

Its proximal extremity is the larger, and is gently curved anconad, 
to meet the corresponding flexure of the brachium to form the elbow- 
joint, the articular surface engaged being quite extensive and vertically 
expanded. The lower, circular, and concave trochlea is the greater sig- 
moid cavity, and is intended for the ulnar tubercle of the humerus. 
Its proximal margin is so produced as to form a strongly defined "ole- 
cranon process", the lower lip of the cavity being the homologne of the 
'-eoronoid process", and is so ieebly developed as to scarcely deserve 
the distinction. In close proximity to the greater sigmoid cavity, above, 
there is another articular surface, quadrilateral in outline, decidedly 
concave from above downwards, much more shallow in the opposite di- 
rection, for the oblique tubercle. Immediately beyond its distal mar- 
gin is a weak and shallow facette for the side of the head of the radius, 
so that the oblique tubercle articulates in a cavity furnished by the 
cupped head of the radius and the larger quadrilateral trochlea of the 
cubitus, the two being almost continuous. 

The outer aspect of this extremity presents simply certain feeble ele- 
vations and depressions for muscles and the ligaments surrounding the 
joint, 

From the proximal extremity, the nearly cylindrical shaft curves 
gently palmad from its inner third only ; after that it takes a compara- 
tively straight course for the wrist. The anconal aspect of the shaft 
presents at the junction of the inner and middle thirds an eliptical nu- 
trient foramen, that enters the bone almost perpendicularly to its long- 
axis. The tubercles for the insertion of the bases of the quills of the 
secondaries, so prominent on this bone in some birds, as in Colaptes, 
seem to be entirely absent. TVe find them barely present in Harporhyn- 
chus, quite strongly marked in Laniua, The carpal extremity of the 



shufeldt.] OSTEOLOGY OF THE EREMOPHILA. 647 

ulna is likewise articular, being vertically cleft and curved downwards. 
Anconad it develops a rough eminence, and above a depression for the 
fan-like expansion of the radius. This end, as in the majority of the 
class, articulates with the three carpal bones and the radius above. 

The humerus measures 2.4 centimetres, the ulna 3 centimetres, and the 
radius 2.7 centimetres; so that when the bones are placed in situ and 
the wing closed, the anti-brachium projects beyond the brachium about 
5 millimetres. The bones of the forearm, though hollow, are apparently 
non-pneumatic, as is the case with the carpals and long bones of the 
manus. 

As in the great majority of the class, the bird- wrist is composed of 
the two free carpals and the os magnum, which is confluent with the 
proximal extremity of the second metacarpal. 

The superior and smaller carpal is the scaphoid, here an irregularly 
shaped bonelet, introduced among the cubitus, the radius, and the conflu- 
ent os magnum, with a distal articular face for the latter and two proximal 
ones for theirochlese of the anti-brachium. Between the scaphoid and the 
cuneiform, the other free and inferior carpal, there exists an interspace, 
where the ulna meets the os magnum. 

The cuneiform has an elongated facet on its outer aspect for the ulna, 
and^two articular processes that grasp the metacarpal below — an arrange- 
ment that admirably meets the action required of the avian wrist. 

The last carpal merely constitutes the trochlear head of the confluent 
metacarpals; by a gentle and backward sweep its general surface is di- 
rected inwards. . 

The composition of the metacarpal bone of this bird does not deviate 
from the general rule, as applied to the class, in any important particu- 
lar. The three long bones comprising it are firmly anchylosed together 
and bear the fingers. The shortest and first metacarpal, obliquely fused 
with the anterior and upper end of the second, supports a free and pointed 
index digit. The second, or medius, supports, first in order below, a 
phalanx peculiar to birds, that is at once recognized by its expanded 
posterior border. It is here deeply concave on its inner surface, which 
concavity is partially divided by a feeble transverse line. 

The blade of this bone is quite thin in some birds, even the general 
surface is sometimes absorbed, leaving nothing but the rounded and 
limital borders, as in Larus delawarensis and others. 

The neck of this bone is but moderately constricted between the blade 
and articular facet for the metacarpal to which it belongs. It bears be- 
low another, and the smallest, phalanx of the hand, a little, free, sharp- 
poiuted and compressed finger, that completes the skeletal bird-arm dis- 
tally, being the ultimate segment. 

The third metacarpal, termed annularis, a slender, ribbon-like bone, 
fast above and below to medius, and extending slightly beyond it, also 
articulates distally with another free phalanx, of the general character 
as the index digit and the ultimate joint of the mid-metacarpal, although 
it is longer than either of them. Measuring along the anterior aspect, 
from the summit of mid-metacarpal to the point of the last phalanx, we 
find the manus in Eremophila to average 2.6 centimetres. 

This, the pectoral limb, as we have endeavored to picture it in this 
Lark, with its brachium, anti-brachium, and pinion in proportionate 
equipoise as to length of segments, with its various bones smooth, 
straight, and devoid of those evidences of being acted upon by powerful 
muscles, would require but a glance from the student of avian skeletol- 
ogy to pronounce it as belonging to a bird possessed of a flight barely 
mediocral in rapidity and power. 



648 GEOLOGICAL SURVEY OF THE TERRITORIES. 

Of the pelvic limb— (PL IV, Figs. 22, 39, 40, 42, 44, and 46).— The inner 
aspect of the upper extremity of the/emw presents the usual globular 
head for articulation with the cotyloid ring of the pelvis. It is nearly 
sessile with the shaft, the neck amounting to almost nil. A shallow and 
inconspicuous excavation occurs on the head for the insertion of the 
ligamentum teres. The articular surface that originates with this hemi- 
spherical protuberance extends outwards over the summit of the bone, 
constantly spreading, until limited by the trochanterian ridge, in a 
plane with the outer aspect of the shaft; it occupies a slightly higher 
level than the head, and it is opposed to the anti-trochanter in the articu- 
lated skeleton. 

Anteriorly the trochanterian ridge and line are quite prominent, ex- 
tending a short distance down the shaft, to be lost on the general surface ; 
posteriorly it projects outwards horizontally from the articular surface, 
over a shallow concavity that is found immediately below, that presents 
at its base a circular foramen that leads to the hollow shaft, and is 
probably the pneumatic foramen, though the femur of this bird does 
not have the appearance of a bone possessed of pneumaticity ; the ori- 
fice, if nutrient, is certainly situated in &n unusual place, though we 
must confess that a careful search over the entire shaft with a powerful 
lens has failed to reveal any other opening. The trochanter minor is not 
represented. 

The shaft, for the greater part of its extent, is cylindrical, and decid- 
edly convex forwards, with a clean superficies, undivided by any inter- 
muscular ridges or lines, or if so, they are very faintly marked. 

The distal extremity of the femur enters largely in to the knee-joint, and 
is more bulky than the proximal extremity of the bone. It is directed 
backwards, and, as usual, is divided by an an tero- posterior shallow in- 
tercondyloid notch, which is continued up the shaft anteriorly, as the 
" rotular channel," soon to disappear into internal and external condyles. 
The larger and lower external condyle is longitudinally cleft posteriorly, 
so as to afford an additional and outer condyloid surface for the head of 
the fibula, with which it articulates. 

A tuberosity is found behind, just above this cleft, and a few others, 
less prominent and situated more internally, are seen on this aspect of 
the bone, in the popliteal fossa. 

The limiting margin of the internal condyle is sharp and distinct. 
The ordinary features, as tuberosities and muscular lines and markings, 
usually sought for at this end of the bone in nearly all birds, are very 
feebly reproduced in our present subject. 

The proximal extremity of the tibia has a very interesting form, due 
to the prominence of the cnemial ridges. These are attached to the head 
of the bone, well above the horizontal articular surface for the condyles of 
the femur. Their superior border is continuous and convex upwards ; 
their inferior borders meet the shaft abruptly, and there terminate. 
Both of these wing-like processes are turned towards the fibular side of 
the bone, the procnemial process being the larger in every respect ; and 
the ectocnemial sometimes is produced downwards into a very sharp 
and needle-like spine, a characteristic of other Oscines. They include 
between them a triangular concave and rather deep recess. The expan- 
sion supporting the superior articular surface projects over the shaft of 
the bone in all directions, being quadrilateral in outline, and having an 
articular facet for the fibula on the outer side, while in the middle of 
the surface above a tuberous spine of the tibia exists, with concavities 
on either side for the condyles of the thigh-bone. 

The shaft is remarkably straight, light, and hollow 7 though apparently 



bhufeldt.] OSTEOLOGY OF THE EREMOPIIILA. 649 

non-pneumatic, no apertures having been discovered to allow the air 
access to the interior. 

A fibular ridge, 4 millimetres long and 1 millimetre deep, is developed 
in the upper third of the shaft, perpendicular to its outer aspect, for the 
lower articulation of that bone. 

Huxley and Gegeubaur maintain that the distal extremity of the 
tibia represents the astragalus among the Class Aves, and there certainly 
seems to be some foundation for this assertion, for if we examine this 
bone in the young of any of the Gallince, as in Centrocercus, we find the 
segment that eventually ossifies with this end of the tibia to be rather 
too extensive for a mere epiphysis, and may represent that tarsal bone. 
This joint is more thoroughly discussed in the osteology of the Tetra- 
oniclce, further on. Without further remark, then, upon this important 
and still unsettled question here, we will observe that in Eremophila, 
and in all birds, the leg-bone terminates distally by two anteriorly placed 
condyles, separated by a well-defined intercondyloid notch. These con- 
dyles, approaching each other behind, diverging in front, are reniform 
in outline and shape, with their convex surfaces downwards. They are 
higher on the shaft anteriorly, and the articular portion is more exten- 
sive. Likewise, anteriorly the shaft is grooved below, to be bridged over 
just above the notch by a narrow bony span, arched outwardly, that holds 
the tendons of the deep extensors in position. 

The inner end of this arch is the higher on the bone, and just above 
it, on the shaft, we find a minute tubercle, that gives attachment to a lig- 
ament that is extended to another tubercle lower down on the shaft, and 
on the opposite side. 

The fibula is the merest apology for a bone, represented only by a 
slender spine on the outer side of the tibia. It has a superior and knob- 
like head, that articulates with the horizontally expanded head of its 
sizable companion ; lower down it meets the fibular ridge, and is firmly 
attached to it by a strong, fibrous, and close-fitting connection. 

Below the ridge the fibula is continued, hair-like in dimensions, to meet 
the tibia below the middle of the shaft, to become thoroughly and indis- 
tinguishably confluent with it. 

The patella (PL IV, Fig. 22) is a free bone, and is found in the tendon 
of the quadriceps extensor. It is compressed antero-posteriorly, with 
an elliptical base above. From the points representing the vertices of 
the major axis of this ellipse, bounding lines pass, to meet broadly con- 
cave below. The anterior surface, limited by these boundaries, is con- 
vex outwards; the posterior surface, slightly concave, is divided by a 
vertical ridge into two unequal parts, the outer of which is the greater. 

The femur averages 2 centimetres in length, the tibia 3.2 centime- 
tres, and the bone now to be described as the tarso-metatarsus nearly 
2.3 centimetres. 

The metatarsals of the second, third, and fourth toes, and certain tar- 
sals at the upper extremity of the bone, coalescing, form the segment, 
tarso-metatarsus, next in order below the tibia, with which it articulates. 

The articular surface of its summit is so arranged as to accommodate 
itself to the condyles of the tibia, consisting essentially of an inner and 
outer antero-posterior facet, and a prominent spine on the anterior mar- 
gin, that accurately fits in the intercondyloid notch of the bone above. 

On the posterior aspect of the bone above we find the il calcaneal" 
process, here approaching a right parallelopiped in form, being vertically 
pierced by four minute cylindrical canals, two next the shaft and two 
parallel with them and above. They are for flexor tendons, which pass 
through them. The shaft is straight, subcylindrical, and hollow, ex- 



650 GEOLOGICAL SURVEY OF THE TERRITORIES. 

paneling below for the trochlear for the phalanges. For its upper half 
and posteriorly, ranging below the calcaneal process, it develops a sharp 
vertical crest, that gradually subsides below. 

The anterior aspect of the shaft is faintly grooved longitudinally, 
and where it dies out below, just above the notch between the third and 
fourth terminal trochlea?, we observe a minute perforating foramen for 
the anterior tibial artery. Upon the inner margin of the shaft below 
there is the well-marked though shallow facet for the os metatarsale acces- 
sorium. This diminutive bone is, as usual, slung to the tarso-metatar- 
sus by a ligament, articulating beyond with the hallux. It represents 
the first metatarsal, and has all the appearances of one of the larger-sized 
phalangeal segments, divided obliquely through the shaft, with the cut 
surface closed in and forming the articular surface for the tarso-meta- 
tarsus. Its position, in situ, is figured in PI. IY, Fig. 44. The lower and 
expanded end of the tarso-metatarsus, bearing the trochlea? of the re- 
maining phalanges, is further conspicuous for the marked manner in 
which the bone is compressed antero-posteriorly, causing the trochlear 
ends to be placed side by side, transversely. The middle one is the 
largest and grooved entirely round, the one for the second toe being 
slightly the higher and bent a little outwards ; finally, the fourth is the 
smallest. Slit-like spaces among these " processes n completely divide 
them. 

The joints of the toes are arranged upon the most common plan, and, 
we believe, upon the general rule for all Oscines ; i. e., the hallux pos- 
sesses two phalanges, second toe three, third toe four, and the outer and 
last toe five. 

These joints are not impressed with anything particularly remarkable, 
beyond what is found in them among the class generally. Their verti- 
cally cleft and anterior extremities articulate with the joint beyond, 
which is diminished in size and articulates in like manner with the next 
anterior segment. 

The claws are grooved laterally, and show a process at their proximal 
and lower aspects. 

A glance at PI. IV, Fig. 22, will be sufficient to satisfy ourselves that 
the great length of the claw of the hind toe sometimes seen in Uremo- 
phila, and always characteristic, is due almost entirely to the growth of 
the horny theca that encases it, and not to the length of the osseous 
claw. 

In the figure just referred to, the hallux, with the first metatarsal, has 
been drawn backwards in the skeleton, not only to show the os meta- 
tarsale accessorium, but also a sesamoid, of no mean size, that is found 
on its outer side, an ossicle that betrays its possessor and declares the 
habit he has of spending a good share of his time upon the ground. 



652 GEOLOGICAL SURVEY OF THE TEEEITOEIES. 



PLATE IV.* 

Fig. 22. Skeleton of Erernopliila alpestris. 

Fig. 23. Sacrum and pelvis from above. 

Fig. 24. The sternum from below. 

Fig. 25. The skull from above. 

Fig. 26. The skull from below. 

Fig. 27. The sternum from above. 

Fig. 28. Sacrum and pelvis from below. 

Fig. 29. Lower mandible from above. 

Fig. 30. The clavicular fnrculum from in front. 

Fig. 31. Left humerus, anconal aspect. 

Fig. 32. Left scapula and coracoid, internal aspect. 

Fig. 33. Scapular extremity of clavicle. 

Fig. 34. Left scapula and coracoid, external aspect, showing extent of glenoid 
cavity. 

Fig. 35. The thirteenth cervical vetebra, showing first pair of free pleurapophyses. 

Fig. 36. Left humerus, palmar aspect. 

Fig. 37. Hyoid arch from below. 

Fig. 38. Anterior view of sternum, first dorsal vertebra, with its movable pleura- 
pophyses and hsemapophyses, in situ. 

Fig. 39. Eight tarso-metatarsus, anterior aspect. 

Fig. 40. Eight femur, anterior asr>ect. 

Fig. 41. Sclerotals, right eye. 

Fig. 42. Eight tibia and fibula, anterior aspect. 

Fig. 43. Eight ulna, anconal aspect. 

Fig. 44. Eight foot, with a portion of the podotheca removed to show tlie os meta- 
tar^ale accessorium, in situ. 

Fig. 46. Eight femur, posterior aspect. 

*The figures on this plate are numbered in continuation with the author's plates and figures to his 
. Memoir on the Osteology of Speotyto cunicularia hypogcea. 



LL. US GEOL SURV. VOL. VI 



PLATE IV. 



FiS 24 



Fi#. 27. 




Big JO. 



OSTEOLOGY OF EREMOPHILA ALPESTRIS. 



OSTEOLOGY OF THE NORTH AMERICAN TETRAONIDH. 



By R. W. Shufeldt, M. D., 

Captain, Medical Department, United States Army. 



The representatives of the Gallinaceous order of birds in the North 
American fauna are referred to four families, the Cracidce, the Melea- 
gridce, the Tetraonidce, and the Perdicidw. Members of the family Or a- 
cidce or the Curassows, are all American birds, being distributed over 
the continent from the Rio Grande southward. 

The latter group has been divided by Messrs. Sclater and Salvin into 
three subfamilies, the Cracinw, the Penelopince, and the Oreophasinw. Of 
the fifty or more species making up these subfamilies, at the present 
writing, but one form has been taken within the limits of the United 
States, this being the Ortalis vetula maccalli, the Ghachalaca of the 
Texan s and Mexicans. The subfamily Penelopince, referred to above, 
has been divided into seven genera, of which Ortalis is the last, and 
the one to which our only North American species belongs. Every 
attempt of the author to secure the skeleton of this interesting bird 
has thus far failed, and as he has had no personal experience with 
Ortalis in its native haunts, the reader is referred to the standard works 
upon ornithology for descriptions of this species. In Meleagridw, the 
second family enumerated above, we have but one genus, Meleagris, 
containing the Turkeys, well-known fowls peculiar to North America, 
and of which there are two species, or rather a species, the western 
form, Meleagris gallopavo, and the eastern variety, or subspecies, Melea- 
gris gallopavo americana. 1 

Although the writer has had the opportunity of comparing several 
skeletons and parts of skeletons of both of these forms with the Te'tra- 
onidw, it must be understood that in recording his observations it has 
only been with the view of calling the reader's attention to the osteo- 
logical similarities and differences, en passant. Professor Huxley, in his 
studies of this group, has left but little to be desired in the way of 
osteological descriptions, clearly pointing out differential characteris- 
tics existing -between the Turkeys and the Numididw, or the Guinea 
Hens. It is the third family, the Tetraonidw, in our enumeration that 
has claimed the greatest share of our labor, and to which the title of 
this paper has been awarded in consequence. We have before us a 
complete collection of the skeletons of the members of this group, which 
is unfortunately more than we can say for the partridges, although 
of the latter family we have, we believe, a sufficient number to observe 
their general characteristics. 

The family Tetraonidw includes the Grouse, of which there are six 
genera in North America, containing sixteen species, embracing ten 
varieties. Of these we shall give a synopsis further on, showing their 
principal external characters, as set forth by American ornithologists, 

*A second species, If. ocellata, is found in portions of Yucatan, Honduras, and 
Guatemala. g53 



G54 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



and also from the author's observations, both in the field and from speci- 
mens in his own collection. 

Among gallinaceous fowls the Grouse are known by certain char- 
acters, such as 
the narrow and 
naked supra-pal- 
petral processes; 
by the feathers 
fiWiug the exter- 
nal nasal fossae ; 
by the cervical 
and inflatable air- 
sacs of Cupidonia 
and others; and 
finally by the 
feathered tarsi, 
the naked toes, 
with the peculiar 
horny processes, 
fringing their 
edges In Lago- 
to the very claws, 




VAVtt' 



Ccntrocercus urophasianus. 



pus the feathers of the tarsi are dense, and carried 

this genus thus forming an exception to the last observation. 

In the arrangement of the genera we adopt the high authority of the 
authors of the History of North American Birds (A Hist. N. A. Birds, 
Boston, 1874, Baird, Brewer, and Eidgway). In this work we find the 
following divisions given us, with some few modifications, which the 
reader will readily recognize : 



1. Genus Canace. (American 
Wood Grouse.) 



Family Tetraonidae. j 
(Grouse.) 



2. Genus Bonasa. 
Grouse ) 



3. Genus Lagopus. 
migans. ) 



(Buffed 



(The Ptar 



Genus Cupidonia. 
nated Grouse.) 



(Pin- 



r 



Subgenus Canace. 



Subgenus Dendra- 
gapus. 



fl- 



3. 



Genus Pedicecetes. (Sharp 
tailed Grouse.) 

Genus Centrocercus 

Cock; Cock of the Plains.) 



Species: 
1. Canace canadensis. 
(Spruce Grouse.) 
Canace canadensis frank- 
lini. (Franklin's Grouse.) 
Canace obscura. (Dusky 
Grouse.) 

Canace obscura fuliginosa. 
( Oregon Dusky Grouse. ) 
Canace obscura richard- 
soni. (Richardson's Dusky 
[ Grouse.) 
Species : 

1. Bonasa umbella. (Ruffed Grouse.) 

2. Bonasa umbella nmbelloides. (Gray Ruffed 
Grouse.) 

3. Bonasa umbella sabinii. (Oregon Ruffed 
Grouse.) 

Species : 

1. Lagopus albus. (The Willow Ptarmigan.) 

2. Lagopus rupestris. (Rock Ptarmigan.) 
,3. Lagopus leucurus. (White-tailed Ptarmigan.) 

Species : 

1. Cupidonia cupido. (Prairie Chicken.) 

2. Cupidonia cupido pallidi cincta. (Lesser Prairie 
Chicken. ) 

Species : 
1. Pedicecetes pbasianellus. (Northern Sharp- 
tailed Grouse.) 
Pedicecetes pbasianellus Columbian as. (Com- 
mon Sharp-tailed Grouse.) 
(Sagei Species: 

I 1. Centrocercus uropbasianus. (Sage Cock.) 



I 2 ' 



For the two subgenera of Canace, the principal differences seem to 
be, that the first, Canace, differs from the second, Dendragapus, in that it 
is of smaller size ; has sixteen tail-feathers to twenty of the latter, and 
also Canace has no inflatable air-sac at the side of the neck, which the 
subgenus Dendragapus has. Otherwise the varieties of each subgenus 



SHUFELDT.] 



OSTEOLOGY OF THE TETRAONHLE. 



655 



seem to have arisen mainly, as so many varieties doubtless have, from 
climatic influences, due to their inhabiting various geographical areas. 

Canace canadensis is found ranging from northern United States to 
the Arctic seas; west nearly to the Rocky Mountains. C. canadensis 
franUini from northern Rocky Mountains, near United States bound- 
ary, and west to the Coast Range. Canace obscura ranges in the Rocky 
Mountains to the south of South Pass and Sierra Nevada, north to 
Oregon and south to San Francisco Mountains, New Mexico, and finally 
Canace obscura fuliginosa, northwest coast region, from Oregon to Sitka. 
The latter two varieties intergrade at their limits. (Habitat, from Hist. 
N. A. Birds, Baird, Brewer, aud Ridgway, 1874.) 

The writer has had the pleasure of hunting the Dusky Grouse at 
various times and localities in the mountainous "districts of the Territory 
of Wyoming. On one occasion, in the early autumn of 1877, I found 

myself heading a _^ '_=_'_=. 

file of several -^=^~z^ '^^B ^tKB$=S 0^= ^ 

Sioux Indians, 
winding my way 
over a well-beaten 
game-path, along 
the side of one of 
the highest peaks 
of the Big Horn 
Mountains. My 
companions were 
armed with the 
well-known car- 
bine, and I had my 
double fowling- 
piece, a weapon 
that none of them 
had ever seen be- 
fore and were evi- 
dently quite curious to see the use and effect of. Suddenly we emerged 
into one of those delightful little open spaces that we occasionally find 
occurring even on the sides of the most rugged ascents. It was devoid 
of trees and carpeted with the greenest of verdure. Hardly had we 




Canace obscura. 



entered this area when away went a 



magnificent 



"Blue Grouse" 



from his hiding place and cover. It was the first bird of the kind that 
I had ever seen alive, and the effect could hardly have been less de- 
moralizing than the impression produced upon the renowned Wallace, 
when the first Bird of Paradise dazzled his vision in the jungles of New 
Guinea. Suffice it to say, that, in a twinkling, this vigorous old Grouse 
was entirely out of my reach, and was whirling up the mountain-side 
through the darkpine woods that covered it. 

But my game was not alone in a spot so inviting, for a step or two 
more started two other fine old males of the same bevy. By this time 
self-possession had been entirely recovered, and before this pair could 
reach the confines of the open space in which we were, each had been 
overtaken by the contents of one of the barrels of my breech-loader 
with fatal effect. Before the echoes of the double report had died away 
among the rocky canons, my Indians had these beautiful birds in their 
hands and were closely examining them, apx)arently looking for some 
immense wound that the bore of my gun certainly seemed capable of 
inflicting. The Indian has, we know, an aggravating way of not show- 
ing his wonderment even on the most startling occasions, and it has 



656 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



always been a source of intense gratification to me if I could by any 
means whatever so surprise one of these stolid sods of the forest as to 
induce him to elevate his eyebrows, be it ever so little. On this par- 
ticular occasion the position was mine. I simply shrugged my shoul- 
ders, giving them to understand that it was an extremely simple mat- 
ter, and left them to make what they could out of it. 




Canace obscura. 



All of thebirds of this species taken in the Big Horn Mountains were, to 
the best of my recollection, the form recognized as G. obscura richardsoni 
I do not remember an exception. This is not the case, however, when 
we come to the Laramie foot-hills about the country and less elevated 
peaks lying to the southward and westward of Fort Fetterman. It was 
in this vicinity and near the top of one of the highest hills to the east- 
ward of Casper Mountain that the writer, in the depth of winter, with 
the thermometer many degrees below zero and the snow knee-deep, 
found small bevies of the type bird C. obscura, and succeeded in taking 
several specimens. They seemed to occur more frequently in those hills 
where the immense herds of elk resorted, and perhaps this was due to 
the fact that these animals bared the ground of snow in many places, 
and thus exposed winter berries and other foods that might not other- 
wise have been added to the bill of fare of these Grouse. Centroccrcus 
urophasianus occurs throughout the Northwest, frequenting, almost with 
out exception, the arid plains wherever the sage-brush thrives, the buds 
and leaves of which form its principal food. 

Thus far this genus has contained but the one species just alluded to, 
and although this Grouse occurs over an immense area of country, there 
is not an ornithologist that we are aware of, that has ever detected 
sufficient differences in specimens taken in widely- separated districts 
that would guarantee the establishment of varieties. The author for 



SHUFELDT. 



OSTEOLOGY OF THE TETRAONID.E. 



657 



several seasons had the opportunity to study these birds in the very 
heart of their native hauuts ; observed them under all manner of cir- 
cumstances, both young and old; but the bird and its habits are so well 
known at the present writing that personal experiences will be almost 
superfluous here. 

For the species and varieties of Bonasa, we find that the type, B. um- 
bella, is found throughout the eastern districts, it being the Partridge of 
New England and the Pheasant of the South. This form intergrades 
with the first variety that we find as we proceed westward; that is, B. 
umbel! a umbelloides, a geographical race found in the Rocky Mountains 
and British America. The last variety, B. umbella sabinii, the Oregon 
Grouse, is the Pacific coast form. The best authorities allow that the 
varieties of the type of this Grouse owe their differences in coloration to 
geographical modifications. 

The extent of this intergradation is well described in the History of 
North American Birds, already cited. After presenting a synopsis of 
the varieties, it reads as follows : 

The above synopsis is intended to present in the simplest form the characteristic feat- 
ures of the three definite races of this exceedingly variable species, as exhibited in 
a light-rusty rufous-tailed form of the Atlantic States, a pale, gray, ashy-tailed form 
of the Rocky Mount- 
ains of the United - =-lr - --^^f^^^^-^^^^ff^-^.-^^ ^^- — - _ 
States and British -±|||j ^;~ ~~^S=_ 
America, and a dark- ^Jf^' S?v° .""-'^j ^----^. 
rusty rufous - tailed 
form of the Northwest 
coast region. These 
three, when based on 
specimens from the re- 
gions where their char- 
acters are most exag- 
gerated and uniform, 
appear sufficiently 
distinct ; but when we 
find that specimens 
from the New England 
States have the rufous 
bodies of umbellus, 
and gray tails of um- 
bellgides, and continue 
to see that the transi- 
tion between any two 
of the three forms is 
gradual with the local- 
ity, we are unavoid- 
ably led to the conclu- Bonasa umbella. 
sion that they are merely geographical modifications of one species. 

Pedicecetes presents us with two forms, already given in our enumera- 
tion of species and varieties ; the first, P. phasianellus is found pretty 
generally throughout British America and to the northward of our 
boundary. It also occurs in Alaska. P. phasianellus columbianus sup- 
plants this species, being a variety of it, and is found in the prairie re- 
gions west of the Mississippi River, and has even been taken as far east 
as Chicago. 

In Wyoming, I found them during the spring and winter months in 
large numbers along the banks of the Platte River and the streams 
and creeks that empty into it. During the summer months, however, 
they desert these localities and resort to the hills and mountains, where 
many of them breed. 

As in the genus just alluded to, Cupidonia has its type and a variety. 
42 H 




G5 8 GEOLOGICAL SURVEY OF THE TEKKITORIES. 

Cupidonia cupido being found in the more central portions of the United 
States,. and at one time, we are well aware, was plentiful all along the 
Atlantic Coast, particularly to the northward. G. cupido pallid ianeta is 
a variety occurring from Texas to Nebraska, generally lighter in color, 
and decidedly smaller. This is one of the most interesting members of 
the family Tetraonida\ and descriptive ornithologists have delighted in 
repsenting us in their works with the engaging history of the famed 
prairie hen. 

There are three species of ptarmigan found in Xorth America, as al- 
ready shown in our table given above. Lagopus albus is confined to 
sub-Arctic and Arctic America, occurring more or less abundantly across 
the continent. L. rupestris is also an Arctic bird, while our last species, 
L. leucurus, is not only found iu these sub-polar regions but inhabits the 
mountainous districts of the West as far south as Xew ALexico. 

I have not had the pleasure of hunting this beautiful Grouse, never 
having been in sections where it was found, so that in the following 
partial synopsis of external characters and appearances of the Tetraonidce, 
I am, as far as this species is concerned, indebted entirely to the authors 
of A History of Xorth American Birds, from which valuable work I have 
borrowed the necessary data differentiating this genus from others of 
the group. 

GEXEKA OF THE TETRAOXID.E. 

Caxace. — Xasal fossae occupying barely half the culnien. An extensible bare space 
on either side of neck. Xo cervical ruffs. Tail broad and rounded at outer angles. 
Toes without feathers, which occur on the basal membranes between them and the 
entire tarsi. The varieties C. canadensis and C. franklini are distinguishable in that 
the former has a rounded tail that terminates with a lightish brown band, and the 
upper tail coverts have light ash emarginations. whereas tbe latter bird has nearly 
ii square Tail, entirely black, only occasionally showing a light colored tippiug, and 
the superior tail-coverts have extensive vxhite margins. 

Boxasa. — Crested, cervical ruffs of black and particularly soft and glossy feathers. 
Tail broad and nearly square. Tarsi bare below. Anterior scutella? arranged in two 
rows, approaching the Partridges iu this as in many osteological characters. 

B. umbella umbellmdes. — Pale : slaty-gray the prevailing shade. (Cones.) 

B. umbella sauinii. — Dark : chestnut-brown tbe prevailing shade. (Cones.) 

Lagopus. — No cervical ruffs. Tail but slightly rounded. Tarsi and toes thickly 
feathered to the very claws. Species become white during winter season. 

L. albus. — Bill very stout. " Bill as high as the distance from The nasal groove to its 
tip. Tail always black, narrowly tipped with white; wing (except upper coverts) 
pure white." 

L. rupestris. — u Bill slender : disTance from the nasal grove to tip (-35) greater than 
height at base (.27). In summer the feathers of back black, bar. 1 i distinctly with 
yellowish-brown and tipped with white. In winter white, the tail black : the male 
with a black bar from bill through the eye. Size considerably less than that of 
L. albus." 

L. Jeucurus. — " Entirely pure white, including the tail." 

CrPiDOXiA. — Nasal fossae less than one-third the length of the culmen. Sides of neck 
ornamented with long and sharp outstanding tufts, composed of thick black feathers. 
Tail shorter in comparison than any other Grouse. Feathers extend to lower end of 
tarsi. Head slightly crested. Inflatable air-sacs below cervical tufts. Osteologically 
Cupidonia and Pedicecetes are nearly related and opposed to all the other forms or ge- 
nera of the Teiraonida?. As in the following genus. Cupidonia has one well-derined va- 
riety, C. cupido paUidieineta : this form is alluded to in the History of Xorth American 
Birds, above cited, in these words: u In its relations with the C. cupido this race bears 
a direct analogy to Pedicecetes eolumbianus, as compared with P. phasianeVu.s. and to 
Ortyx texana, as distinguished from 0. virginiana. Thus in a much less development 
of the tarsal feathers it agrees with the southern Pedicecetes, while in paler, grayer 
colors, and smaller size, it is like the southwestern Ortyx." 

Pedicecetes. — Xasal fossa? less than half the length of the culmen. Cannot be said to 
be crested, but has the habit of raising the feathers of the crown under the same circum- 
stances that the crested forms elevate their crests. Have seen specimens that certainly 
seemed to possess rudimentary cervical tufts. The mid-pair of tail-featbers produced 
beyond the short and graduated tail proper, their ends being truncate. Tarsi feathered 



SHUFELDT.] 



OSTEOLOGY OF THE TETRAONID^. 



G59 



to the i ops. lu P. pliasianellus the toes are hidden by the feathers and the coloration 
of the two forms differs materially. 

Centrocercus urophasianus. — Nasal fossae nearly two-thirds the culmen. Large, in- 
flatable air-sacs. Not crested. Feathers of throat with spiny shafts, and those of the 
body with large after shafts. Tail long, graduated, and of twenty acuminate feathers. 
Tarsi covered with feathers, which extend over the basal webs. 

As we do not enter especially into the subject of the osteology of the 
family Perdicidce, we will merely present the reader here with a synopsis 
of the North American forms and their habitats. 



GENUS. 



Family Perdicidce. 
(The Partridges.) 



© 



Ortyx . 



Ororlyx . 



Lophortyx 



SPECIES. 



HABITAT. 



Ortyx virginiana. (Ameri-C Eastern United States and to 
can Partridge.) } the plains west. 

O. virqiniana floridana. \ CT „ Tir i a 
( Florida Partridge. ) \ L lorma - 

O. virginiana texana. (Tex- ( Texas and certain localities to 
an Partridge.) \ the northward. 



Or ortyx picta. 
ridge. ) 



(Plumed Part- 



Northern coast region of Cali- 
fornia, Washington Territory, 
and Oregon. 



Orortyx picta 
(Plumed Quail.) 



nlnmifpra (Southern California. A rac 
piumijera. l that simply differ8 slightly i 

( coloration. 



race 
in 



' Lophortyx californica. 
fornia Quail.) 

1 Lophortyx gambeli. 
hel's Quail.) 



(Cali- 



C Valleys to the foot-hills of the 

\ Pacific region. 

( Colorado Valley of the United 

(Gam- j States, north to Southern 

Utah, and east to Western 



< 



I 



Texas. (Hist, of N. A. Birds. ) 






(Scaled C Texss, New Mexico, and Ari- 
l zona. 



(Mas- 



New Mexico, Arizona, and Tex- 
as, and to the southward. 



To this list, my friend Dr. Coues has added Coturniv dactylisonans, 
the common Migratory Quail of Europe, a bird that at different times 
has been imported and set at liberty in various parts of the Union — in 
New England for one locality — and it is at present supposed that this 
Quail will become a bird of the country, as Passer domesticus has. 

I learu also from Mr. Lucien M. Turner, lately returned from Alaska, 
that he has been so fortunate as to find a new race of Lagopus that will 
be duly described in his report. 

The anatomical peculiarities of the order Gallinw has been the favorite 
theme of many an able writer, and we find Huxley, Owen, Gegenbaur, 
Parker, Coues, and others, in their several works, dwelling largely upon 
the osteology of these birds, ably exposing the observed characteristics 
of structure both by pen and pencil; but, as far as our knowledge ex- 
tends, no one has as yet devoted himself to the production of a paper 
devoted exclusively to the osteology of the North American Grouse, 
such as the writer here proposes to undertake with every hope of suc- 
cess, aided as he has been by the kind assistance of many friends in 
different parts of the Union, in sending him valuable material in the 
way of representatives of the Family. 

In this monograph we will omit, as we have in former ones now pub- 
lished, any detailed description of the osseous elements of the ear, or the 
respiratory tube, small sesamoids, or such tendons as may ossify in the 
extremities. The hyoid as an arch of one of the cranial vertebrae evi- 
dently is not included in this category, and will in consequence receive 
the attention it undoubtedly deserves in its proper place. 

The study of the bones entering into the cranial vertebrae has been 
initiated at a stage in the chick's life a few days after birth, and their 



660 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



relation to each other and their development carried np to the adnlt 
bird. It will be observed after a glance at the writer's drawings in 
Plate V that he has chosen the young of that grand old prairie-loving 
Grouse, Centrocercus urophasianus, as an example of the growth of the 
sknll from the time above referred to in the Tetraonince. 

In this plate the first three figures show respectively the sknll of the 
young of the Sage Cock a few days after the parent has led it from the 
nest: 47 from above, 48 lateral view, with mandible, and 49 from be- 
low, the mandible removed. 

Fig. 50 shows the bird in August of the same year, and -Fig. 51 the 

disarticulated skull of the same, whereas in the next plate we observe 

the skull of an old cock of the same species, that has, no doubt, trod the 

prairie for many a season. (Fig. 52.) 

In these birds the greatest amount of difference exists in point of size 

among the sexes and 
in individuals of vari- 
^ ous ages of the same 
M sex 5 so we naturally 
find a corresponding 
amount of difference 
in the sizes of their 
crania. 

Fig. 52 is the skull 
of an exceptionally 
large adult, $ , chosen 
from a bevy of sev- 
eral hundred others, 
with a view of afford- 
ing the reader the op- 
portunity of seeing 
the proportions this 
Grouse may attain, 
as far as this part of 
its skeleton is con- 
cerned. This peculi- 
arity seems to be con- 
fined to Centrocercus, and does not obtain with the other varieties, they 
seemingly arriving at maturity of growth at a much earlier period of 
their existence. Canace ouscura may form an exception to these re- 
marks, but it is certain that it is not by any means so striking a char- 
acteristic in this bird. Another interesting point to be observed here, 
that no doubt has forced itself upon the reader since his inspection of 
the plates already introduced, is the unusual length of time that the 
original bony segments of this Grouse's head retain their individuality 
over others of the class. This is indeed so, and in birds of one or two 
years of age, if we exclude the epencephalic arch of the occipital ver- 
tebra, it is not an unusual occurrence to be able to distinguish all the 
sutural boundaries among the remaining elements, and these appear to 
be persistent when applied to the nasals and the premaxillary bone of 
very old birds. We are all well aware that this rule holds good in the 
common barn-yard fowl. 

Students of the works of that eminent anatomist and observer, Owen, 
will remember that in his Comparative Anatomy and Physiology of 
Vertebrates he seems to accuse the Struthionidcu alone of this singular 
feature, or at least u those birds in which the power of flight is abro- 
gated." Now, such of my readers as have had the opportunity of ob- 




Bonasa umbella. 



shufeldt.] OSTEOLOGY OF THE TETRAONID^. 661 

serving the flight of the " Cock of the Plains," after he has once been 
induced to take wing, will agree that there is anything save an abroga- 
tion of that avian privilege. 

Crania of the North American Tetraonince being placed on the hori- 
zontal plane, as described in ray monograph on the osteology of Ere- 
mophila alpestris (Bull. U. S. Geol. and Geogr. Surv. of the Ters., vol. vi., 
No. 1), we observe that their equilibrium is moderately stable, the ante- 
rior bearing point being the tip of the superior mandible, and the two 
posterior bearing points being the external facets upon the tympanies. 
The angles of the foramina magna average 70° while the centrum of 
the parietal vertebra is the chief bone of what here must be the basi- 
cranii, and is found to be nearly in the horizontal plane ; the neural 
arch of the occipital vertebra being, as a whole, gently convex out- 
wards and lying in nearly the same plane with the foramina magna. 

The Skull. 1 — So distinct do we find the hsemal arch of the first cranial 
or occipital vertebra, and fulfilling such a diverse end, with its ap- 
pendage the pectoral limb, in birds generally, that its description will 
be undertaken further on under the subject of the "scapular arch," and 
our attention be engaged at this point only with the neural or epen- 
cephalic arch of this segment of the cranium. 

The primoidal elements of this, the superior arch of the vertebra in 
question are seen to a greater or less extent in situ in the young and 
"bird of the year" of Centrocercus in Plate V, Figs. 47-50, and in the 
disarticulated skull of the same, Fig. 51, as so, eo, bo, and po, lettering 
respectively the essential elements " superoccipital," "exoccipital" (the 
X)arial bone and counterpart of this segment being intentionally omitted, 
as are the duplicates of other segments), "basioccipital," 2 and the con- 
nately developed process, "paroccipital" of the neurapophyses. 

In Sage Cocks the size of those figured in Plate V, Figs. 50 and 51, 
we find the neutral spine of the first vertebra, so, to be a light, spongy 
bone, one and a half centimeters wide by about one-half of a centimeter 
deep — covered with a thin layer of compact substance. Its upper border 
displays in the median line a demi-lozenged si i aped notch that when 
the bone meets the parietals, which latter have their posterior and inner 
corners deficient, forms in many birds of this age a fontanelle. In 
younger individuals this diamond-shaped vacuity is always present, the 

1 The author's plates and figures illustrating this paper are numbered in continua- 
tion with others of his published monographs. 

The reader is kindly referred to the loot-Lote under "Skull" on page 594 of this 
volume, in my monograph upon the osteology of Speolyto cunicularia hypogwa, where 
he will find the author's remarks upon the theory of the four cranial vertebrae, and the 
reasons why they were adopted in certain of his papers. These remarks apply with 
equal force to the present article. 

2 The terms here used for the elements superoccipital, exoccipital, and basioccipital, 
seem to be more or less universally adopted and applied by anatomical writers. Pro- 
fessor Parker and Gegenbaur term the first-mentioned element the supraoccipital 
rather than the snperoccipial, as here given. The former of these comparative anat- 
omists says, in his Morphology of the Skull (Lond., 1877, p. 241) in regard to these seg- 
ments as they are found in the chick of the common fowl the second day after hatch- 
ing: "The extensive occipital plane, swelling backwards above, is largely ossified, 
although there are considerable chondrous tracts remaining. The basioccipital ex- 
tends into the occipital condyle, and it is considerably unci -rflcored by the basitem- 
poral plate in front. (Pig. 63.) The exoccipitals ossify the lower half of the sides of 
the foramen magnum. They are very irregular in shape, extending considerably into 
the ear-cartilage. Th y are perforated by the vagus and hypoglossal nerves. The 
supraoccipital centers have coalesced almost completely to form a large bone bound- 
ing the upper half of the foramen magnum, which is pointed above. Superiorly, the 
margin of the bone is curved like a fan, and abuts on the parietals." The supraoccip- 
ital centers, according to Professor Parker, are two in number, developing side by side. 
{Op. cit., page 235.) 



662 GEOLOGICAL SURVEY OF THE TERRITORIES. 

anterior fontanelle being formed in them in a similar manner, though 
narrower and longer, between the frontals and parietals. The lower 
border of the superoccipital presents a smooth, angular depression, that 
in the articulated vertebra goes to complete the superior third of the 
foramen magnum. 

The lateral bodies of this bone are cellular masses with several aper- 
tures opening forwards and outwards, the mastoids closing them in, in 
the completed cranium. On its outer surface near the inferior angles 
we observe two, one on either side, grooved foramina, leading upwards 
and inwards, to open into the lateral sinuses on the inner surface of the 
segment, nearer together. As age advances these canals contract, but 
still exist throughout life. 

The basioccipital segment, bo, also is largely cancellous in structure, 
wedge-shaped, having at its apex a long, rounded tubercle curving 
outwards and backwards, overhanging a slight depression beneath it. 
This tubercle in the complete vertebra forms the middle two-thirds of 
the occipital condyle, which, in the adult, is found below the foramen 
magnum, sessile, uniform in outline, with the rounded border below, and 
all indications of its original division into three parts obliterated. 

The neurapophyses of this vertebra, termed the "exoocipitals" (Plate 
V, Fig. 51, eo) 1 are each nearly as large as the neural spine ; on their inner 
borders they present for examination the deeply-rounded margins to 
complete the foramen magnum, and immediately beneath, the minute 
tubercle jutting out that lends its assistance on either side to form the 
condyle of the occiput. 

The outer angles, quadrate in outline, deflected slightly downwards, 
are the transverse processes of the vertebra, the "paroccipitals." The 
precondyloid foramina are also to be observed here, with one still more 
external, belonging to the group from which the eighth nerve makes its 
way from the cranium. The internal aspect of an exoccipital is a mass 
of open, irregular cells, that are closed in when this segment is approxi- 
mated with the mastoid, superoccipital, and the " petrosal ", l (Fig. 51,1), 
that odd-shaped and spongy bonelet which constitutes the capsule of 
the organ of hearing — and which has a foramen on its inner and smooth 
surface for the passage of the auditory nerve — forming', also, by a bend- 
ing forwards of a part of this surface, and aided by the basi sphenoid, 
the floor of the mesencephalic fossa on either side, while externally it 
shares in forming the entrance from without to the otocrane. 

With the exception of the petrosal, the elements thus far described, 
when duly articulated, form the neural arch of the occipital vertebra, as 
already intimated above. The basioccipital, the centrum of this verte- 
bra, by its larger extremity and the exoccipitals with the connate 
diapophyses articulate with the basi-sphenoid in the basi cranii below; 
the latter, with the superoccipital, meet the parietals and mastoids above 
and laterally. In old birds every trace, both sutural and otherwise, 
becomes completely obliterated as the osseous amalgamation pro- 
gresses, though throughout the group a well-defined "superior line" 
limiting muscular attachment, indicates very nearly the terminating 
borders above, and sometimes, as in Centrocercus, a fainter indication 
exists in the vicinity of the union among the interested bones below. 
On either side of the condyle to its outer aspect we observe in a slight 
depression a group of usually four foramina — two external opening into 

1 Quite recently Dr. Coues has, in an admirable paper, presented us with a review of 
the literature bearing upon the so-called "temporal bone" of human anatomy, and as 
in this paper the true relations and composition of the " petrossal" are so clearly s t 
forth, and they express the views now generally accepted, that it gives me pleasure 



shufeldt.] OSTEOLOGY OF THE TETKAONID^. 663 

the otocrane, one into the cranial cavity, aud one leading through the 
basi-sphenoid to the base of the " sella turcica'" at the carotid opening; 
they transmit principally the eighth nerve and the internal jugular aud 
branches. 

In some of the very old individuals of the Tetraoninw quite a striking 
characteristic presents itself in the capaciousness of the opening to the 
otocrane, produced by a thin, wing-like expansion, recurved forwards, 
formed by the outgrowing aud union of the centrum of the second 
vertebra and the diapophysis of the first. This feature is not particu- 
larly noticeable in the Ortygince, nor in Lagopus, Cupidonia, and Bonasa, 
still less so in the Sharp-tailed Grouse, among the Tetraonince, but quite 
marked in old males, especially in Canace and Centrocercus (Figs. 52, 
74, 88, and 89) . No very decided differences exist among the Grouse 
with regard to the foramen magnum and the occipital condyle ; the 
former is universally of good size for its owner, subciicular, and with- 
out any encroachments upon its margins beyond the condyle. This lat- 
ter, always sessile, reniformin contour, occupies its usual position below 
the foramen, with its long axis placed horizontally. In all the Grouse, 
save Canace and Centroeercus, it slightly invades the marginal periphery 
of the great foramen of the occiput, and in all the excepted genera is 
more or less shortened transversely. 

The second cranial segment constitutes the parietal vertebra, and its 
elements are shown in the same plate, Fig. 51, where indicators pass 
through its neural and haemal arches, P. V. and P. V: P. Y is the mes- 
encephalic arch, constituted in the complete cranium by the bones P, 
the parietal or neural spine, when linked with its fellow ; a. s., the alis- 
phenoids, the neurapophyses ; m. s., the mastoids, the diapophyses ; and 

to quote the following from him. In summing up the various parts of the temporal 
bone, he says: 

The following is a list of these morphological elements : 

1. Squamosal. A membrane bone with which in mammals the maxilla is connected by the malar and 

the mandible is directly articulated. The ' ' squamous portion " of the temporal. 

2. Tympanic. Accessory to the organ of hearing, forming the meatus auditorius externns. The 

"auditory process." 
3 Pro otic f Primitive otic elements together forming the periotic or petro-mastoid ; the oto- 

' Eoi otic J crane or bony capsule of the organ of hearingj»inclosing the bony labyrinth, 

k rwvsl+i, ,-!+;„ 1 developing the mastoid cells, and practically constituting the "petrous" and 

5. upistn-otic { , , mastoid „ parts of the temporal, 

f 6. Malleus, the proximal end of the mandibular 

6. Mallens. C Ossicula aditus or phonophori j arch 

7. Incas. < in man devoted to audition, v?- Incas, the proximal end of the hyoidean arch. 

8. Stapes. I but | 8. iStapes, connecting fenestra ovalis with the 

{ hyoidean arch. 

Q TvrrmaTin lwal f ( Tymp'ano-hyal not recognized in human anat- 

in i£EPwni Y \ Elements of the hyoidean arch. \ omy. 
iu. sstyio-nyaL £ (Stylo-hyal, the "styloid process." 

This does not include the os orbiculare, a minute ossification occurring at the junc- 
tion of the iucus with the stapes, and in man for a short time separate. 

Still further on this author states : " The periotic develops from amass of cartilage 
situated in the basis cranii between the occipital and sphenoid from three centers of 
ossification, which, however speedily and completely they may coalesce, as they do 
in man before birth, represent as many distinct bones, one or more of which may re- 
main separate in many animals. These are the prootic, the epiotic, and the opisthotic. 
The first of these is anterior and in special relation with the corresponding vertical 
semi-circular canal. The second is superior and external. The third is posterior and 
inferior in relation with the posterior vertical semi-circular canal. Their confluence 
completes a bony periotic capsule, inclosing the labyrinth or cavity of the inner ear. 
This is the triune periotic bone; with its mastoid developments the still only triune 
petro-mastoid bone; with its tympanic annex the otocrane, or skull of the ear, contain- 
ing the essential parts of the organ of hearing. The periotic proper corresponds 
closely enough with the " petrous portion of the temporal" of human anatomy — the 
"petrosal bone/'' as it is sometimes called. (The Nature of the Human Temporal 
Bone, Am, Jour, Otology, vol. iv, Jan., 1882, pp. 25-2D.) 



664 GEOLOGICAL SYRVEY OF THE TERRITORIES. 

b. *., the basi-sphenoid. tlie centrum of the vertebra. The haemal arch 
we see in the •• hyoid,"' which here shares the same fate of its neighbor 
in the occipital vertebra, insomuch as it is ununited to the superior arch 
by either osseous connection or by articulation, for in all living birds 
the hyoid. the well-known bony support of the tongue, depends en- 
tirely upon its muscular and ligamentous connections to retain its rela- 
tions with the cranium. The manner in which the disjoined neural spine 
of the parietal vertebra goes to form the posterior fontanelle in the half- 
grown bird has already been sufficiently dwelt upon. The bone P, as 
detached in an individual of this age, is quadrilateral in outline, ex- 
cessively spongy and light, owing to the paucity of compact substance 
over quite a large share of diploic tissue, which is chiefly deposited in 
a protuberance on its inner table, which protuberance, in union with 
the fellow of the opposite side of the complete cranial vault, forms two 
concave surfaces out of the remainder of the superficies, essential por- 
tions of the ep- and prosencephalic fossa?. 

Superficially, these elements are smooth and convex, and in the adult, 
after consolidation, exhibit some faint evidence of a parietal eminence 
on either side — more marked elevations, however, occurring in the spine 
of the vertebra beyond, immediately anterior to the suture termed in 
Anthropotomv the "coronal." With the exceptions of the tympano- 
mastoidal articulation and the connections between the mastoids and 
petrosals, the majority of the articulations of this vertebra in the mid- 
aged bird may be classed among the variety known and described in 
works upon human anatomy as the "squamosal" — the parietals being 
beveled above to accommodate themselves to the frontals. 

The alisphenoids are separated from each other mesially by nearly 
half a centimeter: above they meet the frontals, below the basi- sphe- 
noid, and laterally the mastoids — the lower and outer angles almost 
reaching the cup shaped articulation tor the tympanies. This segment 
seems to ossify from its borders towards the center, leaving a foramen 
that is eventually closed in. Touching this point, Professor Parker 
says, when treating of the development of tbe skull of the Common 
Fowl, " From the hinder half of the basisphenoidal region a considerable 
elongated alisplienoidal lamina arises [a. s. Fig. 59), passing outwards 
and backwards on eitner side in the cranial floor, and ascending a little 
into the side wall; coalescing at its extremity with the periotic mass, but 
leaving an elongated space unfilled by cartilage between the two tracts. 
The cranial surface of the alisphenoid conforms closely with the concave 
curvature of the hinder part of the cranium (see Fig. 62)." (Morphol- 
ogy of the Skull, Lond. 1877, p. 230). And further on, "A membra- 
nous fontanelle (a. s.f.) arises in its center; and both in front and be- 
hind this distinct ossific centers appear in the cartilage; these after- 
wards unite to form one alisphenoid bone." (Ibid p. 236). 

Professor Huxley, in treating of the development of the cranium, says 
of this bone — 

"In front of the auditory capsules and of the exit of the third divis- 
ion of the fifth nerve, a center of ossification may appear on each side 
and give rise to the alisplienoid, which normally becomes united below 
with the basisphenoid." (The Anat. of Vert. Animals, p. 24.) 

This bone, on its mid and lower border, presents for examination the 
half of the "foramen ovale/" which is completed by meeting the centrum 
of the vertebra. It is for the transmission of the Trigeminal nerve into 
the orbital cavity. Laterally there is developed a quadrate apophysis 
(the parapophysis of the vertebra?), which joins with a similar, subse- 
quently scale-like process coming from the mastoid, resulting in a fora- 



shufeldt.] OSTEOLOGY OF THE TETRAONID^E. 665 

men, cordate in outline above, elliptical below, between them, giving 
passage to the fibres of the temporal muscle that is markedly charac- 
teristic of the Tetraonidcv. Below the point of union this sphenotic 
process, as this latter apophysis has been termed by authors, is triangu- 
lar, with its apex pointing forwards and downwards, flat, with its inner 
surface looking forwards, upwards, and inwards. Internally, the alis- 
phenoid is deeply concave. (Plate VI, Fig. 52, and other skulls illus- 
trating this paper.) 

The external appearance of the mastcid 1 is well shown in Plate V, Fig. 
50, and as m. s., Fig. 51. Internally the half-cells observed close in by 
the aid of similar excavations in the segments of the occipital vertebra, 
the acoustic capsule; and a double-concave surface assists in forming 
cranial fossse. 

We now come to examine the ornithic characters of one of the most 
interesting segments of the bird-skull, the centrum of the parietal 
vertebra, well termed by most ornithotomists and general anatomists as 
the basi-sphenoid. At an early date in the life of the chick (Centrocer- 
cus and others) this bone becomes confluent with the centrum of the 
frontal vertebra be- 
yond; this conflu- 
ence takes place, if 
we may be allowed 
to differ with such 
high authority a s 
Owen, who makes 
the rather sweeping 
assertion '-that the 
pit for the pituitary 
body marks the 
boun d a ry " ( C o m p . 
Anat. and Phys. of 

the Vert VOl. ii. T). Cupidonia cupido. 

45) in the following manner, and the sutural trace is yet discernible in 
young birds of the Family under discussion (Plate V, Fig. 51). The 
jjre-sphenoid lies beneath a tuberous process projecting anteriorly from 
the latter bone, reaching nearly as lar back as the carotid foramina. 
The combined bones, the centra of the two mid-cranial vertebras, thus 
constitute the compound bone basi-pre-sphenoid of comparative anato- 
mists. 

Viewed from above, we discover, proceeding from before backwards, 
in the median line, 1. The upper aspect of the apophysis just mentioned, 
and immediately to its rear che deep " sella turcica" with the osseous 
caials of the carotids opening into one foramen at its base; 2. Two 
smooth surfaces, one on either side and a little laterally, for the optic 
chiasma to rest upon ; 3. Another surface still more posteriorly for the 
mesencephalic fossa, being perforated by diminutive parial foramina; 
4. A roughened open space for the articulation, with the head of the 
wedge-like basi-occipital. Anteriorly, and at the same time laterally, 
broad and uneven borders for the alisphenoids, with their smooth groove- 
lets of the foramen ovale, while back of these again, on either posterior 
angle, a concave wing-like expansion, the terminations of the Eustachian 

1 The bone here called mastoid is the squamosal of the majority of authors, as Huxley, 
Parker, and others. It corresponds with the hone that hears the same name in the 
works of Professor Owen. The hone squamosal of this latter author, and used in the 
same connection by myself in this monograph, is the quadrato-jugal of Huxley and 
others. 




666 GEOLOGICAL SURVEY OE THE TERRITORIES. 

tubes, that add to the parietes of the entrance of the otocrane. Below 
and superiorly, at the base of the junction of the two boDes. we find the 
carotid foramina, with a depression between them mesially, and still 
lower dowu. slightly protected by an attenuated offshoot from beneath, 
the separate apertures of the anterior and buccal entrances of the Eus- 
tachian tubes. 

The remaining surface, unbroken in character, extending posteriorly, 
goes to complete the basi cranii. The coaptation of the elements form- 
ing the neural arch of the parietal vertebra is shown in Plate V, Figs. 
47-50, their amalgamation in the adult in Plate VI, Fig. 52. 

The chief importance of the haemal arch of this vertebra depends 
upon the bony support it affords the tongue. In a fine specimen of an 
adult Lagopus leueurus, kindly presented me by Mr. Robert B. MeLeod, 
then residing in Leadville. Colo.. w*» find the following characteristics 
presented to us for examination, and they extend with little deviation 
to all the members of the family. The hyoid arch 1 consists of. in the 
specimen under consideration, seven bones. The continent ceratohyals 
and glossohyal. which latter is largely completed anteriorly by cartilage. 
form one segment: the ceratohyals diverge from each other smartly 
behind, and at their point of meeting afford the facette for the trans- 
verse trochlea surface on the basi-hyal. This last bone, the second in 
order, measures half a centimeter in length, being enlarged at both ex- 
tremities, flattened from above downwards, the anterior end being fash- 
ioned to fulfill the purpose already mentioned, while the posterior and 
larger extremity presents two faeettes, looking backwards and outwards, 
to articulate with the hypobranchial elements of the thyro-hyals. The 
third segment also meets this compound articulation at this point, a 
short urohyal. it. too. being completed at its posterior extremity by car- 
tilage. The hypo- and cerato-bratiehial elements oi'the thyro-hyals make 
angles with each other and curve upwards in conformity with the basi- 
cranii. 

The sub-cylindrical hypo-branchials are one and a half centimeters 
long, and connected with the posterior elements by quite along and in- 
tervening -piece of cartilage of the same caliber: the smaller cerato- 
branchials also taper off behind with the same material. 

This arch in the letraonidcelong remains almost entirely cartilaginous. 
the hypo-branchial elements alone being composed of bone, and a bird 
must be of quite an advanced age before he can boast of a complete 
osseous framework as forming a component part of his lingual apparatus. 

The neural arch of the third cranial vertebra now to be defined is the 
prosencephalic — its lneinal arch the "mandibular."' as its tueinapophy- 
sis constitutes the lower jaw, termed " mandible.*' in avian skeletology. 

The fusing of the centrum of this segment with the basi-sphenoid has 
already been elucidated: the rostrum thus lormed is gently inclined 
upwards and forwards, grooved along its entire superior aspect, tapering 
to a sharp point anteriorly to receive the connate prefrontals in the bony 
gutter at its distal third. Beneath it displays towards its base the 

1 In speaking of this compound bone, Professor Gegenbaur Tells us : '"Two pairs of 
arches can be made out in birds. The rudimentary nrst arch fuses to form the so- 
called entoglossal bone (Fig. '259, 2). posteriorly to which lies the true body of the 
hyoid. The second arch, however, is well developed, and gives rise to the coruna 
(4. 5), which are formed of two large pieces, which generally curve backwards behind 
the skull, without being directly connected with it. Behind the copula there is the 
remnant of a second one. which forms the hyoid process (3)." (Elements of Comp. 
Anat.. Lond.. 1878, p. 472.) In the description of the hyovd in this paper the cerato- 
hyals correspond to the entoglossal bone, as referred to in the above quotation from 
Gegenbaur. and the copula of that author to the basi-hyal, his hyoid process being 
the urohval. 



shufeldt.] OSTEOLOGY OF THE TETEAONID^. 667 

parial facets for the pterygoids and beyond the rounded surface for the 
palatine articulation. 

It will be remembered that in the first edition of this monograph the 
writer announced the fact that he had failed to discover the orbito-sphe- 
noids (considered as the neurapophyses of the arch) i. e., they were not 
found as bones that were produced from separate points of ossification. 
Since that time, more than a year ago, I have not had the opportunity 
to look more thoroughly into the mattter, but the series of my skulls 
of the common fowl and those of Centroeercus show the spaces these 
bones occupy in the mature birds to be completely filled in, the posterior 
orbital walls being more or less complete, a fact familiar to all of us, but 
in birds of the year and still younger specimens large vacuities exist 
where these bones should be; moreover, I have in my possession a 
skull of the common fowl in which the basi-sphenoid sends up on either 
side two delicate bony sprouts that subsequently complete the periphery 
of the circular foramen for the oeuloinotorial nerve and the optic. Pro- 
fessor Owen seems to think that these are the centres of ossification for the 
orbito sphenoids, for this anatomist tells us that "the bones. 10 (orbito- 
sphenoids), of the third neural arch coalesce with each other, and the cen- 
trum below, protect a smaller proportion of the prosencephalon than in the 
Crocodile, but maintain their neurapophysial relation to it and to the op- 
tic nerves, below the exit of which they begin to ossify." ( Anat. of Verts. 
Vol. II, p. 46.) Huxley, in his Anatomy of Yertebrated Animals, page 22, 
says, "In front of, or above, the exits of the optic nerves the orbitosphe- 
noidal ossifications may appear and unite below with the pre-sphenoid." 
Professor Parker is even more decided, for this writer informs us that 
"Above the optic foramen, wedged in between the alisphenoid and the 
interorbital septum, a four-sided bone has arisen in membrane on each 
side, and there is a smaller pair in front of and above the larger, helping 
to fill in the fenestra} left unoccupied by the orbital plates of thefrontals. 
These are the anterior and posterior orbitosphenoids (o. s. Fig. 66.) The 
anterior half of the interorbital septum is ossified, the mesethmoid en- 
croaching on the anterior margin of the interorbital fenestra." (Morph. 
of the Skull, young fowls up to nine months old, page 249.) 

This is v r ery clear, and as Professor Parker is a very careful observer 
and devoted himself particularly to the subject in question, no doubt 
his observation is correct, and some more fortunate observer than my- 
self will some day detect these ossifications forthe orbito -sphenoids in our 
Tetraonidce ; for since they occur in the common fowl, it seems only 
natural that we should look for them in the Gallince generally. This 
accounts for the fact that in my drawing in Plate V , Fig. 51, the po- 
sition of an orbito-sphenoid was simply shown by dotted lines, and 
marked os.; in this same figure Fr. is the " frontal," ps. the prefrontal 
or centrum of the i 7 ertebra, and x the usual site forthe postfrontal — this 
exogenous element, the diapophysis of the vertebra is not here found, 
its position being occupied by a depressed roughened surface for the 
squamous articulation of the mastoid. We have never personally ex- 
amined any bird in our avifauna where this bone is seen independent. 
Descriptive ornithotomists, in their studies upon the skulls of Eheidce and 
Struthionidw, give the presence of this process as occurring free. 

The neural spine of the frontal vertebra follows the example of the 
parietal in being completely bifidated in the younger specimens. As a 
whole it is perhaps the largest segment in the bird-skull — certainly as 
far as our Grouse and Partridges are concerned. Either half of its 
spine presents, projecting anteriorly from the middle, a flattened process, 
directed gently forwards, downwards, and outwards; that at its extrem- 



668 GEOLOGICAL SURVEY OF THE TERRITORIES. 

ity is marked above by quite an extensive surface for one of the nasals, 
and below by another, against which the head of the ethmoid abuts. 
The concave surface below this process and the remaining- hinder moiety 
form the vault of the orbit. Another scale-like projection is thrown 
out posteriorly, deeply concave within, correspondingly convex without, 
to shield the prosen cephalic lobes — the bones being joined. Huxley 
terms the pleurapophysis of the haemal arch of this segment the " quad- 
rate^ — the u os quadratum" of the older anatomists. Owen defined it 
as the tympanic, 1 it being the hoinologue of a bone of the vertebral skull 
generally — it was the os carre, in birds, in the writings of the eminent 
Cuvier. The tympanic bones here, as in all saorops da, are free ossicles. 
connecting the articular ends of the lower jaw with the skull. They arc 
in the Tetraonidce, symmetrical and well proportioned, not exhibiting any 
marked peculiarities or deviations from a common type. The mastoidal 
and orbital arms are about of a length and calibre, the first being rather 
the larger, and is surmounted by a hemispherical articulating head for 
the cup on the lower border of the mastoid. The neck below the pro- 
cesses is moderately constricted before it expands to become the ''man- 
dibular" end, that has beneath its tranverse elliptical facet outwardly 
the intervening notch, and then the inner and smaller one, all for artic- 
ulation with the mandible. The bone has likewise a surface to articu- 
late with the pterygoid below the orbital process, and is always pneu- 
matic. From the outer aspect of the mandibular extremity it sup- 
ports its two appendages, the bony styles, termed ''squamosal'" and 
malar — the first by a diminutive ball-and-socket joint articulated in 
the usual manner. The malar, as we know, is the mid-style of the 
infraorbital bar — the maxillary completing the connection anteriorly, 
and although upon superficial inspection of this striking ornithic feature 
of the lateral aspect of the skull, it seems to be firmly united in its schin- 
dylesial articulation, it simply requires ordinary maceration in the adult 
of any of the Grouse or Quails to have the three styles separate from 
each other and from their tympanic and intermaxillary connections. 

The lower jaws of the Tetraonince are singularly alike in all their 
characteristics throughout the sub-family. The single bone is devel- 
oped in the usual way by confiuency of the "articular," "surangular," 
" angular," and •'spleniar' elements posteriorly, and the outer moiety by 
the "dentary" element, the haemal spine. (Plate V. See explanation 
of plates for the above-described bones.) 

The maiidible in the adult has a gentle and increasing curvature 
downwards from the interangular vacuity forwards. The curvatures 

1 There can scarcely be a reasonable doubt left in the minds of comparative anato- 
mists that the question is now settled and the fact generally accented that the bone 
termed tympanic and so used by Professor Owen in his writings, is the representative 
of the malleus of the mammal a. It is the ''quadrate bone" of all modern writers, 
I believe without exception, and articulates with the skull, the quadrat o-jugal and th_ 
pterygoid and the lower jaw on either side. Professor Huxley alludes to its develop- 
ment in the following words: ''In the sauropsidan embryo a rod of cartilage occupies 
the first viseral arch on each side and meets its fellow in the middle line. The rod 
becomes jointed, and the part on the distal side of the joint is converted into Meckel's 
cartilage, while that on the proximal side of the jont is modeled into the rudiment 
of the quadrate bone, which is invariably in its earliest state cartilaginous. Soon, 
however, the quadrate cartilage ossifies, and a centre of ossification appears in that 
part of Meckel's cartilage which articulates with the quadiatutn. This gives rise to 
the articular element of the mandible. All the other constituents of the lower jaw 
are developed in the fibrous tissue which surrounds the rest of Meckel's cartilage, 
which structure either persists throughout life or disappears." (On the Kepreseut atives 
of the Malleus and the Incus of the Mammalian and other Vertebrata. Proc. Zool. Soc. 
Loud., 1869, page 401.) 



SHUFELDT.] 



OSTEOLOGY OF THE TETRAONID.E. 



669 



at the extremities of the syrnphysisial suture are both parabolic, the 
inner being the more open of the two. The interangular fenestra is a 
flattened ellipse, which has distiuct sutural traces leading from it, indi- 
cating the borders of some of the original bits of bone of which it is 
composed. 

The "coranoids" are but feebly developed and the articular ends not 
far below them; these latter have the usual pneumatic ioramen at the 
ends of their in - 

Va '^^^ 17044- $ 



pointed and blunt 
extremities, and 
sharp recurved pro- 
cesses behind, in a 
line with the rami of 
the jaw, which apo- 
physes long remain 
in cartilage in imma- 
ture birds. (Plate 



X, 



Fig. 



71.) 




On the lateral as- 
pect of the bone, two 
muscular lines lead 
away from the coro- 
n o i d a 1 elevations. 
These last two men- 
tioned features are 
universally charac- 
teristic of the Tetra- 
onidw ; they are 
strongly marked in 
Lag opus. (Plate Cupidonia cupido. 

XIII, Fig. 88.) Minute foramina are found above and below near the 
dentary margins, and two quite prominent, one beyond the ramal fen- 
estra on the inner surface of the jaw; still another just anterior to a 
small tubercle below the coronoids on the same aspect. The inferior 
ramal borders are smooth and rounded, as are the under surfaces of 
the articular ends where they originate in nearly the same plane. 
"" The divergence of the ramal limbs of the mandible in Ortyginm is 
greater, owing to the greater width of the skull when compared with its 
length. 

In some fine specimens of Lopliortyx californica, generously furnished 
us by Mr. Charles A. Allen, of Xicasio, Marin County, California, we 
note the striking departure from the mandible in the Grouse in the 
absence of the interangular vacuity — this feature obtains, however, in 
the common Virginia Partridge and others. The deflection of the rami 
anteriorly is greater in these birds also, or at least more sudden, and 
so prominent are these ramal borders that the inner sides, towards the 
posterior ends, are conveited into true fossae. 

Those interesting osseous and diminutive oblong relates, the sclerotals, 
present in all of the class, we believe without exception, are found here 
occupying their usual position. (Plate Y, Fig. 51, 2, aid Plate X, Fig. 
75, in Cupidonia.) They number from thirteen to eighteen or twenty, 
and their function is so well known that it will not be dwelt upon 
here. They differ principally in the amount of tenacity with which they 
retain their normal relation after prolonged maceration. Cupidonia 
holds a high place here, and the fact seems to be due to the greater 
overlapping of the edges of these little affairs and the toughness, or 



670 GEOLOGICAL SURVEY OF THE TERRITORIES. 

perchance the thickness, of the internal and external sclerotic coats 
that cover them. Lately we saw in the case of Sayornis nigricans where 
these platelets were apparently confluent ; no such condition ever occurs 
in the Grouse or Partridges. 

The "lacrynial" (Plate Y, Pig. 51, 3 and other figured skulls), is 
found on the anterior margin of the frontal, enjoying a free harmonial 
articulation that encroaches slightly on the nasal border. Each is a 
squamous, cordate lamella, with its larger end nearer the orbital cavity ; 
this completes the bone in young birds, but in mature individuals it 
sends down a curved and delicate style with its point directed outwards, 
that encircles and gives support to the lacrymal duct on its passage to 
the rhinal cavity (Centrocercus). 

We now come to examine into the last of the cranial vertebras, and, 
in the family under consideration, the one most modified. It is the 
" nasal," and its neural arch the " rhinencephalic," the haemal, the 
" maxillary." 

In the letraonidce its centrum, the " vomer,' 7 is missing. 1 Whether 
this be due to the foreshortened skull of the Grouse, with its long sphe- 
noidal rostrum rendering any further extension superfluous, we cannot 
say. In the lengthened skull of any of the Anatidoe, where such a bone 
is imperatively called for, as a sub-interspinal partition, it is invariably 
present and unusually prominent (Plate V, Fig. 51, vr. vomer, is merely 
outlined to indicate its position in other birds). 

The neurapophyses of the arch are found in the connate prefrontals, 
the bone called "ethmoid" in androtomy. It here, in the young bird, 
is lodged in the outer third of the groove on the pre split noid, rises 
columnar, sub-compressed laterally, leaning forwards at a gentle angle 
to expand above in a trihedral summit for the support of the frontals, 
nasals, and intermaxillary, a short process being projected backwards 
for the former. The posterior aspect of the column develops as the bird 
grows, the interorbital septum, reaching to, and perhaps aiding in, the 
formation of the exogenous orbito-sphenoids. 

The nasals, or the divided neural spine of the arch, are squamous 

1 Since making the above statement I have been able to examine a large mass of 
material from all imaginable sources, and have come to the final conclusion that the 
vomer occurs in the entire group of gallinaceous birds of America. Many things may 
have led me to believe at the time that this bone was missing, as stated in the text : 
In the first place, it is an extremely delicate and freely articulated one in the Teiraon- 
idcu, and in the second place the doubt still harasses my mind as to whether there 
may not be instances where this vomer does not ossify until very late in the life of the 
bird, and all of the fresh specimens I examined may have been of this character. The 
foundation for this assertion lies in the fact, that quite recently I examined, with 
great care, numbers of market specimens of Bonasa and Cupidonia, in which I failed 
to discover this bonelet, while in others it was easily found. The specimens exam- 
ined were all apparently birds that had attained maturity, and displayed no external 
characters by which they could be separated from each other. 

The plates accompanying this edition were not submitted to me in time to add this 
bone to such figures in which it would show, so that the vomer does not appear in 
any of them. In Plate V, fig. 51, where it is showu in dotted outline as rr., it should 
be beyond the ethmoid, Pf. and shaded like the other bones indicating its presence. 
A good figure, giving its proper position, is presented to us by Huxley in his Anatomy 
of the Vertebrated Animals, page 283 (figure 82 of the common fowl). This writer 
tells us "the vomers vary more than almost any other bones of the sknll. They under- 
lie and embrace the inferior edge of the etlimo-presphenoidal region of the basis cranii, 
and, in all birds in which they are distinctly developed, except the Ostrich, they are 
connected behind with the palatine bones. In most birds they early unite into a 
single bone; but they long remain distinct in some (Joracomorphw, and seem to be al- 
ways separate in the Woodpeckers." 

I have found it in numerous adult specimens of the common fowl, but it is a rare 
coincidence to ever find it in the crania of museum collections. I have in my section 
at the United States National Museum, the "Darwin types" of the skulls of the wild 



shufetdt] OSTEOLOGY OF THE TETEAONID^E. 671 

lamella, twisted upon themselves in a manner to conform with the 
superior base of the beak, overlap the frontals, as already defined, are 
separated from each other by the intermaxillary, throwing out below to 
meet this bone a sharp process, thus forming a broad elliptical bound- 
ary limiting the capacious osseous nares. In all adults of this family 
they are easily detached by maceration. These bones are well shown 
in Plate X, Figs. 71 and 73, in the cranium of Gupidonia, from an un- 
usually fine bird sent with a number of others, for which our thanks are 
graciously tendered to Captain Richards Barnett, Medical Department, 
United States Army. It will be observed that the bone becomes so at- 
tenuated in some specimens as to give rise to a foramen, as seen in the 
latter figure. The haemal arch of this vertebra is called the maxillary 
as its lower rib and spine constitute the major share of the superior man- 
dible or maxilla. . The pleura pophyses seen in the palatines are long, rib- 
like bones with their anterior ends much flattened from above down- 
wards, to fit into a fissure on either side made for them in the inter- 
maxillary below the maxillaries. Near their middles they curve moder- 
ately outwards to develop compressed heads at their posterior extremi- 
ties, fitting into a notch in either pterygoid, and concave mesially for 
the rostrum of the basi-sphenoid. 

At their inner thirds they send off thin sheets of bone that curve up- 
wards, barely to touch the ramphosial process of the sphenoid, accom- 
panying it as far as it extends distad, then sloping away on the ribs of 
the bones themselves. The haBmapophysial maxillaries are elements 
that seldom change their ornithic characters, and in the Tetraonidceseem 
to be reduced to their simple typical form — in completing the delicate 
infraorbital bar on the one hand — and just previous to becoming wedged 
into the premaxillary above the palatines, dispatching a bony offshoot 
on either side nearly to meet each o'ther in the palatine fissure on the 
other. 

The remaining pair of bones found at the interior aspect of the bird's 
skull are the pterygoids. In the Grouse they are stumpy, subcompressed 
concerns, with half-twisted shafts, having broad concave surfaces for 
the facets on the rostrum, which are notched distally for the reception 
of the palatines. The articulation with the tympanies is equally exten- 
sive, monopolizing long, narrow facets beneath the orbital processes on 
those bones. 

We have arrived finally at the point in our descriptive skeletology of 
the avian skull, where we have to deal with the anterior and ultimate 

GaUas baukiva and the Horned Fowl of the Azores. They were presented to the Smith- 
sonian Institution by Mr. Teg< tmeier. The vomer is present in neither of these 
crania, although no doubt it may have been originally, and in the drawings made 
from these very crania the vomer has likewise been omitted. (Darwin's Animals and 
Plants under Domestication. Vol. 1, pp. 275, 279, figs. 34,36.) 

Other eminent writers describe its development in the following language: "The 
pre-maxillaries and the rest of the bones of the upper jaw are further advanced in 
development, becoming more solidified, and perfecting the various relations already 
described. A new bone has arisen in the palate, viz, the vomer (v. fig. 65, p. 246), 
which is however but a very small style, lying under the nasal septum, behind the 
points of the maxillo-palatine processes of the maxillaries." (Parker and Bettany, 
Morph. of the Skull. Section 549. Common Fowl, fourth stage.) And again: "The 
slender maxillary (figs. 64, 65) now rises in front into the angle between the descend- 
ing crus of the nasal bone and the hinder edge of the dentary process of the pre- 
maxillary. The maxillo-palatine plates (Mx. p. Fig. 65; are broader and reach nearly 
to the mid-line, being separated partly by the nasal septum and partly by the small 
vomer, which is rounded in front and split for a short distance behind. The forks of 
the vomer (v.) articulate with the iuner and anterior points of the inner plates of the 
palatine bones, which lie side by side mesially, nearly concealing the rostrum." (Ibid. 
sect. 563. Fifth stage. The chick second day after hatching.) 



B72 



II 51 1 SURVEY OF THE TEEBTTOEIES. 



haemal spine, here fulfilling most important functions as the superior 
mandible, as it does throughout the class at large. In the TetraomMa . 
in the vast majority of birds, the intermajFillafif or the "preniaxillary ~ 
.v.vrhors is of moeh stouter material than most other bones of 
the head, its nse being a very obvious reason for this. (Plate V, Fig. 
1 . N. Pf. nra, L mx.) 

Wi _ Hie moderately free fronto-maxillary and pseudo hinge-joint, 
between the out-turned frontals, the culmen of this bone slopes by a 
gently increasing arc to the tip of the beak. This surface is rounded 
and split in two from the enlarged inner extremity to a point over the 
distal border of the nostril; this division lasts during life. The exter- 
nal nasal orifices are unusually large and sub-elliptical in outline. The 
head of the ethmoid shows in very young chicks, but is eventually cov- 
ered bv this bone, which also fills in snuglv the interaasal space (Plate 




d .:-: :■■: ■.'•'..-'.. :: ". z.r.^ v.-/. 



The osseous maxillary tomia are even sharper than when they were 
capped with the horny integumental sheath that the .entire bill wears 
during life ; they are produced backwards on a triangular process of the 
bone below the shaft of the mamillaries, touching them in the Quails. 



TO 



shufeldt.] OSTEOLOGY OF THE TETRAONID^. 67 

A row of minute foramina encircle the beak anteriorly, where it is the 
thickest, though the segment is non -pneumatic. The general surface 
beneath is depressed below the tomial margins, though it is not very 
extensive, as the wide palatine fissure occupies a good part of the space, 
that termiuates anteriorly in a U-shaped curve, opposite the outer nasal 
border. In the Ortigynw the curve of the culmen is more abrupt, 
and the frontals rise above, in some cases even jut over, the pre in axil- 
lary. The nasal apertures are also very large and of a shorter elliptical 
outline; the palatine fissure is likewise narrower in comparison, a few of 
which differences are such as one would naturally look for in a bird of so 
near kin, and whose beak has been more than proportionately curtailed. 

On removing the vault of the cranium in an adult female of Centro- 
cercus, so as to obtain a free view of the brain -case, we discover the 
usual nervous and vascular foramina present at their most common 
sites, but beyond this we are more struck with the feebleness with 
which many of the salient points are developed, as compared with some 
of the other avian groups; we might sum it up by describing it as a 
lack of angularity and depth. It is true the various fossse are well, 
though not strongly divided, the superior median crest is present, but 
not very prominently developed, and the rhinencephalic fossa is barely 
conical. The section shows the greatest amount of deploic tissue to be 
in the basi-sphenoid, and bones of the occiput, where for potent reasons 
such material is most urgently in demand. 

In the study of the crania of the adult Tetraonidce as an entirety we 
find among the most conspicuous features enlisting our interest the un- 
usual number of bones that remain free in them. The skull can be so 
stripped of its outstanding segments that nothing remains save the 
cephalic casket with the interorbital septum. The rhinal chamber is 
strikingly open, due to the great external nasal passages, and all its in- 
ternal structures, as the ethmo-turbinals, internasal septum, and floor 
being formed only in cartilage. ' A pocket existing in the extremity of 
the premaxillary, that fills in with a spongy osseous tissue during life, 
is observed in Centrocercus, which is solid in the Ortygince and Lagopus — 
parial, subcircular pits placed side by side in like locality in Canace 
obscura. 

The orbits are more fortunate in the completeness of their bony in- 
closures — the heavy plate generously extended by the ethmoid to divide 
these cavities very rarely shows any deficiencies. Of all the crania be- 
fore us Canace obscura is the only delinquent in this respect, though no 
doubt this may occur in others. In it quite a vacuity exists near the 
middle of the septum. Anteriorly the prefrontal and frontal throw out 
laterally squamous septa of greater or less completeness, that divide 
these cavities from the common rhinal space. These plates may coa- 
lesce with the processes of the lacrymal, as a rare coincidence, and per- 
chance meet the infraorbital style. The foramina for the passage of 
the optic nerves and the first pair are, as a rule, singularly circular and 
distinct, the minor apertures about them enjoying a like individuality. 
They are noted for their greater size among the Partridges. 

A separate canal is devoted to each olfactory nerve immediately be- 
low the orbital vaults, that usually at its outstart from the cranial end 
has a small opening between it and the one of the opposite side. About 
the entrance to the otocrane we notice principally a rounded, squamous 
plate thrown down from above by the mastoid, that is present in all tbe 
Grouse. Just below and within, this segment also develops a sharp 
spicula of bone, posterior to the tympanic articulation, that evidently 

43 H 



674 GEOLOGICAL BUKVJSV J THE TERRITORIES. 

serves the purpose of keeping this free ossicle in its socket in certain 
movements of the ja « . 

Peculiarities of the floor of this cavity have already been described 
above : in specimens of Cam/nee mmademmis. carefully selected for me by 
Mr. William Brewster, of Cambridge, Mass-, and forwarded to me by 
Prof. J. A Allen, of the Mnseum of Comparative Zoology, also of Cam- 
bridge- to whom my grateful acknowledgments are dre for so many like 
favors, we find, upon viewing the ->ull from below T the elevations or 
convexities due to these ellipsoidal and wing-like formations, reminding 
one of their marked resemblance to the acoustic bulls of the tympanic 
found among the crania of Felidct. 

The author in his various plates and figures believes he has given suf- 
ficient life size, as they all are, views of basal and superior aspects of 
the skulls of these birds, will not enter here into any needless details 
of measurements. The variation in size in this respect in Centroeercmg 
has already been dwelt upon ; it is not nearly so marked in other genera. 
The surfaces of the skulls above have a rough look caused by many 
minute depressions and groovelets, these running out to the margins of 
the orbits cause them in some to be finely serrated. 

The sharp-tailed Grouse is a unique exception to this, it being a bird 
of rather a delicate skull with smooth cranial superficies. AT - >*- 

troeerems possess rather depressed foreheads, apparently due inpre to a 
slight tilting upwards of the superior orbital peripheries. The lateral 
temporal fossa? are shallow and scarcely noticeable, the muscles they 
afford lodgment not being remarkable either for their size or strength. 

Did the writer feel that he had sufficient material before him he would 
gladly devote a few of these pages to the description of some of the 
exceedingly interesting osteological differences existing between the 
domestic and the wild Turkey : but as such facts can only be considered 
reliable, and such differences constant, after the examination of a large 
series of each, such as we have not at present at our hand, we will 
simply speak of a few of the cranial peculiarities as seen in a set of 
skulls of Jidleagrig galloparo and M. gallapazo americawa from the col- 
lection of the Army Medical Museum of Washington. As we might 
expect, the skull of Mdeagris has in it all of the leading points that we 
have attributed to the Tetraomidm generally. The occiput and the for- 
amen magnum are found to be nearly or quite in the vertical plane, the 
skull resting on its bearing points in the horizontal plai:e ; the surface 
above is more or less rough and venated, as in Cmpidoma, The parietals 
rise above the general surface as rounded domes, constituting quite a 
prominent feature in the skull of this bird. A depression occurs in 
the frontal region between the margins of the orbit and posterior to the 
nasals : this is more decided in my specimens of Pavo erigtatmz, a bird 
that possesses a cranium not at all unlike the Turkey. In Meleagris 
we find the lacrymals to be strong and pointed bones directed back- 
wards, articulating principally with the nasals, though the frontals 
usually extend down to meet their posterior borders ; below, their de- 
scending processes are flattened and turned towards the median plane. 
Taken as a whole the lacrymals of the Wild Turkey are quite different 
bones as compared with the same bones as we found them in the Grouse 
and Quails. A very interesting change is seen to take place in these 
bones in Xttmida wteleagris. as this bird attains maturity. We have just 
said that the frontals in the Turkey extended down so as to meet and 
articulate with the posterior borders of these bones on either side ; now. 
the older the bird the more extensive is the meeting of these bot 
this is carried to its maximum condition in XumidOj for in the young of 



shufelpt] OSTEOLOGY OF THE TETRAONLD.E. 675 

this fowl the lacrymals are found to be freely articulated to the outer 
borders of the nasals, the frontals barely making any encroachment upon 
them, but in a very old bird we find the lacrymals actually wedged in 
between the frontals and nasals on either side, and the superior peri- 
phery of each orbit formed by the continuous outer margins of a frontal 
and a lacrymal. The prominent bony crest seen in the median line of 
the skull of the Guinea Hen is formed entirely by an uprising of the 
frontals and increases as this fowl becomes older. 

In the Wild Turkey, the external nasal apertures are large and ellip- 
tical, and the various sutures among the bones of the superior mandi- 
ble, long remain clear and distinct as we found it to be the case in the 
Grouse. On a lateral view of the skull we find the arrangement of 
the sphenotic process the same as in all of the Tetraonidce ; it differs in 
Numida by being single and very stout ; there seems to be no rule gov- 
erning the condition of the orbital septum, for in skulls of Wild Turkeys 
of apparently the same age — in one, large deficiencies will be found in 
this plate while in the other the partition will be entire and quite thick. 

The pterygoids are large and stumpy bones, articulated in precisely 
the same manner as we described them for the Grouse. The vomer is 
short and the spinal chamber in the dry skull capacious and undivided 
by an osseous septum narium. 

The lower jaw is stout and may or may not have the fenestra present 
in its side ; behind it has the sharp, upturned processes so character- 
istic of Gallinaceous birds. The ramal apertures is absent in all of 
my specimens of Pavo and Numida, otherwise the birds have inferior 
maxillae typical of the family to which they belong. 

We have examined skulls of all of the forms of Lagopus occurring 
in America, and discover but trifling differences existing ; in a speci- 
men of L. albus the orbital septum is complete and the cranium of the 
bird is broad across the fronto-maxillary region; a deep median pit ex- 
ists in this region among many of the Quails, less marked in Ortyx, but 
very decided in Cyrtonyx massena, still more in Orortyx picta, the beau- 
tiful plumed Partridge of the Pacific States. In this latter bird we 
find a deep and longitudinal cleft occurring in the median line on the 
superior aspect of the skull, between the orbital margins, that is char- 
acteristic and not found among the other American Quails. 

The skull of the young of Pedicecetes, a few days after it has left the 
nest, differs in no great degree from the skull of the young of Centrocer- 
cus — i. e., in points of ossification and the relation of the bones. 

Of the Vertebral Column. — In discussing the development and peculiar- 
ities of the vertebral column, we will still continue to adhere to Gentro- 
cercus as our model, explicitly stating names of other species when oc- 
casion requires a departure therefrom. 

In examining the atlas and axis as they occur together in the chick a 
few days old, we find that the neurapophyses of the first have as yet 
failed to fairly meet above in the median line ; though they may in 
some instances, as they undoubtedly do, soon touch each other. No os- 
sific centre exists for an atlantal neural spine, as that process is not found 
upon this bone in any of the Tetraonidce. Below the arch the interest- 
ing procedure is progressing in the appropriation of the centrum of this 
segment by the axis. The inferior extremeties of the atlantal neura- 
pophyses have inserted between them a circular ossicle whose plane is in 
the horizontal plane and on a level with the floor of the neural canal of 
the axis. This bonelet eventually becomes the " odontoid process" of the 
second vertebra. At this stage it is a little less than a millimeter in 
diameter, and in the adult occupying the same position remains a sub- 



676 GEOLOGICAL SURVEY OF THE TERRITORIES. 

sessile, and in comparison with the bulk of the bone it is attached to, an 
insignificant tubercle, though unquestionably fulfilling all the impor- 
tant functions required of it. . In less than six weeks the odontoiclal lar- 
ceny is complete, and no trace remains to lead one to suspect how mat- 
ters stood at the earlier date. 

Immediately beneath and a little posterior to the primoidal and dis- 
tinct centrum of the atlas, th< j re is another, and still larger, free ossific 
centre, uniform in outline, concave above, surrounding the primitive 
odontoid apophysis with its long axis lying transversely; behind, and 
in contact with it, are two more very minute and elliptical ones placed 
side by side. The first of these unite with the atlas and latterly form 
the bony ring for the occipital condyle to revolve in, and the surface for 
the odontoid to move upon, and a notched lip of bone that projects from 
it behind, that subsequently develops; the remaining two, behind the 
first ossicle mentioned, form the anterior part of the body of the axis 
that bears the articulating surface for the first vertebra. In the full- 
grown bird the postzygapophyses of the first vertebra projecting well 
to the rear look almost directly inward. They meet the prezygapoph- 
yses of the axis that face in a coutrary direction and a little backwards. 
The articulating facet for the centrum of the axis is subelliptical, convex, 
of some size, and has in the segmented column the inferior and convex 
surface of the odontoid playing just above it, the superior and flat sur- 
face of the latter being confined by intervening and delicate ligament 
forming a part of the floor of the neural canal of this bone. In the axis 
of the adult the anterior part of the bone with the odontoid process, 
that was separately added, projects conspicuously forwards beyond the 
neural arch, and in birds of several months of age it can be distinctly 
discerned where the union was established between neurapophyses and 
centrum. 

In the mature vertebra the neural canal is nearly circular. In the 
center of the bone, above, a knob-like tubercle acts as the neural spine, 
which has mesially and behind a deep pitlet for the insertion of the in- 
terspinous ligament. Anapophysial tubercles are found above the pcxst 
zygapophyses, which latter are of considerable size, concave, and faced 
downwards. The centrum of the axis is subcompressed from side to side 
and supports mesially, just anterior to the second vertebral articulation, 
the first hypapophysis of the series. The first two segments of the ver- 
tebral column are non pneumatic. 

Vertebrae throughout the chain in the young chick invariably show 
the line of union between the centra and neurapophyses, but it is lost 
as soon as the birds come to be two or three months old. At this age, 
however, still very interesting points of development are strikingly visi- 
ble in the third vertebra, which otherwise varies but slightly from the 
same bone in appearance as seen in the column of an old male, such as 
we have before us. 

The neural spine, more compressed than in the axis, is nearer the mid- 
dle of the vertebra, still deeply pitted for the interspinous ligament be- 
hind, and slightly so on its anterior margin . This characteristic becomes 
very faint among the long vertebrae in the middle of the neck, to be more 
strongly reproduced as we approach the dorsals, the posterior depres- 
sion always being by far the best defined. We find anapophysial tuber- 
cles still present in the third vertebra. These also exist throughout the 
cervical series, with more or less clearness ; they form ridge-like lines 
upon tke.elongated segments of the mid-neck. The zygapophysial pro 
cesses in general look upwards and inwards anteriorly, and vice versa be- 
hind — the fourth vertebra having in common with the one we are now 



shufeldt.] OSTEOLOGY OF THE TETRAONIDiE. 677 

describing an ihterzygapophysial bar, lending to these two segments 
that broad and solid appearance well known to ornithotoinists, not pos- 
sessed by any other of the cervicals. The neural canal in the third ver- 
tebra is nearly circular, which is also its form in the ad nit, becoming 
only moderately compressed from above downwards in the last three or 
four cervicals. Regarding the third vertebra from below, we observe 
that the articulating surface of the centrum for the axis to be quite con- 
cave and turned a little downwards. The processes that fall beneath the 
prezygapophyses form what would be a canal with its lateral margins; 
this groove, however, in the "bird of the year" is converted into the ver- 
tebral canal by an independent ossicle being placed over it on either 
side, and, being below the rest, it causes a broad shallow concavity to 
appear mesially and anteriorly. 

These small bones have at the very outstart stumpy apophyses project- 
ing backwards, which are the parapophyses of the vertebra — the pro- 
jections they meet from above being the pleurapophyses, the groove they 
form mesially being the broad termination of the carotid canal. 

The fourth vertebra has the same general appearance of the third, that 
we have just been describing ; it is a little longer, however, and in both 
large pneumatic foramina are found laterally and beneath the diapoph- 
yses. These apertures are found in the vertebral canal in the remain- 
der of the cervicals. Again, in both, the bodies are rather compressed 
from side to side, and it is not until the bird has arrived at maturity that 
the hypapophyses are well seen in these two segments. 

Now, taking up the cervicals from the fifth vertebra, we find certain 
characteristics holding good throughout the series, with certain gradual 
modifications. In the adult the neural spine in the fifth is prominent and 
placed anteriorly ; it slowly subsides to the tenth, where it is more tuber- 
ous, nearer the middle of the bone, and bears evidence of having a 
posterior projection overhanging the depression for the interspiuous 
ligament. This is the type to include the thirteenth, the projection being 
more and more prominent and slightly cleft behind ; in the fourteenth 
and fifteenth it suddenly assumes the broad quadrate spine of the dor- 
sal type. Returning to the fifth vertebra, we note another change in 
the lengthening of the postzygapophyses ; the acme of this modification 
is seen to be in the sixth and seventh vertebrae. From these they gradu- 
ally shorten again, while the anterior ones spread out with the dia- 
pophyses to assume the form of the consolidated ones in mid-dorsal col- 
umn. This arrangement allows lozenge -shaped apertures to exist be- 
tween the segments above, and subelliptical ones laterally, that become 
smaller and more circular above as the postzygapophyses shorten, and 
quite large laterally as they approach the point opposite where the bra- 
chial plexus is thrown off from the myelou. 

In the adult Cock of the Plains we detect beneath, in the fifth ver- 
tebra, well anteriorly, a strongly -developed quadrate hypapophysis. 
This process entirely disappears in the sixth, for in this segment the 
centrum of the bone, anteriorly on either side, just where the parapo- 
physes meet the body mesiad, a tubercle commences to make its appear- 
ance, the apices slightly inclined towards each other. From the sixth to 
the tenth inclusive these apophyses become longer, approach each other 
below, but never meet so long as they have the "carotid canal," which 
they form between them. In the eleventh they seem to have met through- 
out their extent to form a hypapophysis on the exact site they occupy 
in the tenth, the tenth vertebra being the last cervical where there is 
any evidence of the carotid canal ; hence from this method of formation 
Professor Owen is made to say (Comp. Anat. and Phys. of Vertebrates, 



G78 GEOLOGICAL SURVEY OF THE TERRITORIES. 

vol. 11, p. 190), "In the common Fowl each carotid * ■* * enters (ing) 
the canal formed by the hypapophyses." 

In the completed twelfth vertebra of mature birds we find this 
hypapophysis very large, with expanded extremity, and the parapophy- 
sis, on either side, sending down long subsquamous processes. In the 
thirteenth segment of the " bird of tlie year " theparapophyses begin to 
take on a change. This change develops in the adult still a perfect hypa- 
pophysis, but in the younger individual the parapophysial element 
begins to be notched anteriorly, a part favoring the pleurapophysis, a 
part the centrum, so that in the fourteenth vertebra of the adult the 
hypapophysis is still present anteriorly with a tubercle developing on 
either side of it, with the parietes of the vertebral canal very much 
slenderer. In examining this segment in the younger bird we ascertain 
that the original ossicle is now a descending pleurapophysis meeting 
the parapophysis, a delicate and independent process, which, in the fif- 
teenth and last cervical vertebra, constitutes a free rib, while the hypapo- 
physis consists of a mid process and a smaller nodule on either side. 
This beautiful metamorphosis can be thoroughly studied and easily com- 
prehended in the cervical portion of the vertebral column in our Catliar- 
tes aura. 

So that, as a partial recapitulation of the first fifteen segments, we 
find that they make up the " cervical portion" of the column. Their cen- 
tra are universally subeompressed at their middles, they develop in the 
young bird parapophysial projections that eventually produce free ribs 
by the aid of the descending pleurapophyses, and their interarticula- 
tions, as far as their bodies are concerned, bear out the general ornithic 
law of being apparently procoelous on vertical section and opisthocce- 
lous on horizontal section. 

Backwards from the fifteenth the vertebral segments or the links of 
the chain take on a metamorphosis that is characteristic of the Tetraon- 
idcc. It consists in, in all the adults of the genera, a consolidation ot 
the ensuing four vertebrae. The confluent boue thus formed constitutes 
the major part of the dorsal division of the spinal column and invariably 
supports free pleurapophyses (Plate VI, Fig. 5o, Centrocercus, ad. S ). 
In Centrocercus these four vertebrae can easily be distinguished from 
each other until the bird is over a year old, but very soon after this all 
sutural traces are entirely obliterated and we have the segment as rep- 
resented in the plate. 

The neural spines become one long parallelogrammie plate, occasion- 
ally exhibiting a for- 
amen or so at the 
site of the original 
interspinous spaces. 
Its crest is round- 
ed, but has no inde- 
pendent rim. Mus- 
cular fascia attached 
to it posteriorly often 
ossifies, leaving in 
Pedicecetes phasianeiius. the prepared skele- 

ton flattened spicule, on either side, directed backwards. The anterior 
aspect of this bone has all the necessary elements to meet the last free 
vertebra beyond it. The first pair of diapophyses are the shortest, the 
last pair the longest and most raised; these processes are more or less 
bound together by inetapophysial offshoots of variously defined serrate 
margins, that allow iuterdiapophysial vacuities to exist. Below, aud 




shufeldt.] OSTEOLOGY OF THE TETEAONID^E. 679 

just anterior to the bases of the diapophyses, are the four subelliptical 
and concave facets of the capitula of the dorsal pleurapophyses. From 
their upper and posterior points sharp crests run beneath the transverse 
processes to meet the out-turned and cordate facets at their extremities 
for the tubercula of the ribs. At regular intervals, and nearly in a 
right line among the diapophyses, are the elliptical orifices for the 
transmission of the dorsal nerves. 

Pneumatic foramina are found about the bases of the transverse pro- 
cesses, the most anterior one being of some size. 

The centra of these amalgamated vertebra are very much compressed 
from side to side. This compression has its due influence upon the form 
of the neural tube within, while the bone below is produced downwards 
into an excessively thin and sharp crest, which is still further extended 
into four inverted T-shaped hypapophyses of large size. They join each 
other beneath to a greater or less degree in many of the genera. This 
union more frequently takes place between the first and second, and the 
first is always upturned and flanged out laterally, a feature prominently 
reproduced in Lagopus and Bonasa. These winglike side extensions of 
the lower margins of the hypapophyses not infrequently are continued 
on two or three consecutive ones, and are sometimes the widest on the 
second. — Canace. 

Both in the Grouse and Partridges we find a free vertebra inserted 
between the confound bone we have just been discussing and the first 
sacral vertebra. This segment we must consider as belonging to the 
dorsal series, although in Cupidonia and Pedicecetes the upper and distal 
aspects of its diapophyses are more or less moulded to conform with the 
ilia. This bone is also figured in Plate VI, Pig. 55, in conjunction with 
the other dorsals, with which it has all its characteristics in common, 
and such ones superadded as we might expect to find in a vertebra 
naturally disjointed in the middle of the column. It has the longest 
diapophyses of any of the series to which it belongs ; facets for the 
tubercula and capitula of its own free pleurapophyses ; a hypapophysis 
of no mean length that may or may-uot be expanded below. 

Metapophysial spicules on its transverse processes sometimes are so 
far produced as to reach vertebrae before and behind it. 

This segment is likewise pneumatic. 

There is a wonderful vein of regularity running through the dorsal 
pleurapophyses, hsernapophyses, and hsemal^spine or sternum. As to 
the first pleurapophysis, or rather the first pair of these bones, we have 
already described them as they are found in the ultimate cervical. 
There we are aware it never reaches the sternum by the intervention of 
a sternal rib 5 that they are usually found to possess thoroughly devel- 
oped heads and tubercles for the vertebra, their shafts being less flat- 
tened than the dorsal ribs, and only in Lagopus did we discover any 
epipleural appendages. 

As far as our observations extend, the consolidated portion of the 
column of the dorsum has always consisted of four vertebrae, and con- 
sequently we find in this portion four pairs of movable dorsal pleura- 
pophyses and one pair for the free dorsal vertebra, reckoning five alto- 
gether for this region. The first pair of dorsal vertebral ribs terminate 
in free extremities, which are usually in line with the inter-ha3ma-pleura- 
pophysial articulations in old specimens, and as a rule support epi- 
pleural appendages, characteristic of the species to which the rib 
belongs. 

In the Tetraonidce, following the general aphorism of the class, we ob- 
serve a gradual dilatation of the skeleton from the inter- coracoidal aper- 



680 GEOLOGICAL SURVEY OF THE TERRITORIES. 

ture towards the pelvic extremity of the body. To compensate, or rather 
what really gives rise to to this, as far as the bird's osseous structure is 
concerned, among other things, is a lengthening of the diapophyses as 
we proceed in that direction. The ribs also become longer: the xiphoidal 
forks of the sternum flange outwards. So, too, we hnd a corresponding 
shrinking of the tubercula on the dorso-vertebral pleurapophyses, and 
a stretching of the correlative necks, so that the capitula may reach the 
facet intended for them on the vertebra?,. 

The vertebral ribs as seen in the Grouse are pneumatic, the foramina 
being found on the posterior aspect of these hones immediately below 
the tubercles, sheltered by the flattened bodies as they dilate in that 
region. 

As in nearly all birds, these ribs are compressed from within outwards, 
and in Centrocercus are wider below the uncinate processes, contract- 
ing as they meet their sternal ribs. In these birds, too, the epi-pleural 
appendages are firmly knit to the body of the rib — though in Bonasa, La- 
gopiiSj and Canace they may be easily detached, leaving behind them in 
each case a shallow concave facet. In the last dorsal rib this append- 
age is the smallest, and is directed upwards nearly parallel with the body 
of the rib, and sometimes (Centrocercus) the edge of its rib swells out to 
meet its apex above; and this may have been the method by which the 
unique and remarkably striking uncinate appendages were formed in 
Cupidonia, a bird that constitutes itself an exception among the North 
American Grouse in this respect (PI. XI, Figs. 79, 80), where not only 
the vertebral ribs are unusually broad, but their processes much more 
so, being great, odd, quadrate and compressed plates confluent with the 
body of the rib, and only rarely condescending to be weakened by the 
merest apology for a foramen to exist in them, as in Fig. 80. The Sharp- 
tailed Grouse appears to come next in approaching this state of affairs; 
it, roo, invariably developing more or less expanded ribs. The only 
sacral pair of pleurapophyses conform to the general shape of the series 
to which they belong, and never support epi-pleural processes. 

These bony protectors of the thoracic parietes were found to be 
exceedingly delicate in specimens of Bonasa kindly sent me by Leslie 
A. Lee, Esq., of Brunswick, Me. 

Four haemal ribs ascend from the costal borders of the sternum to 
meet the dorsal pleurapophyses in arthrodial articulations above the 
mid -horizontal plane of the bird's body. As is usual, their sternal ends 
are twisted so that the facets are situated transversely, while the upper 
extremities are broad and flat, particularly in Cupidonia and Pedioecetes. 

The first pair are generally in advance of the others, and insert them- 
selves higher up on the costal processes : The pneumatic foramina for 
these bones are to be found just above their transverse facets anteriorly. 
The extremities of the haamapophyses of the sacral ribs meet the pos- 
terior borders of the ha?mal ribs of the last dorsal pair usually about 
the junction of their middle and lower thirds of the bodies of these 
bones. 

The distal or upper extremities of these, the last pair of hsema- 
pophyses, are sometimes very much expanded, and in a specimen of 
Cenlrocercus, 9 , a small bit of bone has been superadded, simulating an 
additional hffimal spine on either side, as if anticipating the descent 
of another pair of sacral ribs to protect this otherwise feebly guarded 
region of its owner's anatomy. 

The short pair of xipboidal processes overlap the ultimate sternal ribs 
of all the Grouse and Quails, on either side, as do the ilia the sacral pleu- 
rapophyses above. 



shufeldt.1 OSTEOLOGY OF THE TETRAONID^E. 681 

The Sternum,m the North American Tertraonidce, is developed from Jive 
points of ossification, and to these it seems to have added, later in life, 
or before the bone becomes one entire piece, an ossific centre at the ex- 
tremity of each of the four lateral xiphoidal prolongations from which 
their subsequently dilated ends are produced. These later are easily to 
be demonstrated in the haemal spine of Centrocercus, in the "bird of the 
year" (Plate VI, Figs. 53 and 56). 

Fig. 53 represents the young of this last-named Grouse a day or so 
old, at which time all live of the primoidal points of development are 
eminently distinct. The "body" of the bone is nearly circular. The 
" keel," of which only the anterior part has as yet ossified, dips well 
down between the tender pectorals; the manubrium, now only in carti- 
lage, has at this date no evidence of the foramen that later joins the 
coracoidal grooves. As to the rest, bands of delicate membranous tissue 
bind them loosely together. The sternum in a bird of several months' 
growth is show T n in Fig. 50. Here the bone is rapidly assuming the 
shape it is destined to retain during life. The body and with it the keel 
is extending by generous deposition of bone tissue at its margins, prin- 
cipally at the mid-xiphoidal prolongation. The manubrium, still in car- 
tilage, we find pierced at its base by the foramen just alluded to, and 
a rim of the same material runs about the anterior border of the lophos- 
teon, Fig. 56, 4, while a rapidly diminishing band also connects the ele- 
ments known at this stage as the plearosteon, *&., 6, and the metosteon, ib. y 
5. In cases where severe maceration is resorted to with this bone, in 
still older specimens, in which the sutures are not suspected, these parts 
will still separate about the original points of ultimate union. 

On the reverse side of the bone shown in Fig. 56 we find that even 
at this stage it is deeply perforated by the pneumatic foramen at a 
point immediately over the carinal ridge. 

In the adult the sternum is highly pneumatic, air having access to it 
through such apertures not only at this point but also in the costal bor- 
ders between the sternal ribs, and by a single foramen in the groove, 
posterior to the manubrial process mesiad. 

In Plate VI, Figs. 52, 54, and 55 are all parts of the skeleton of the 
same bird — an old adult Sage Cock, Centrocercus — of which Fig. 54 is a 
view from below of the sternum. 

It will be seen that it has a length of 14 centimetres, and other meas- 
urements can be easily obtained from it. We have never seen this bone 
any larger, and, as it is, it represents the maximum size the b genial 
spine attains among North American Grouse. The bone is shown in other 
plates also, and their owners can be ascertained if the reader will kindly 
refer to the description given opposite each plate. 

The unique outline of the sternum of the Gallinw has long been known, 
many authors having both figured and described it, and we will say here 
that in the Tetraonidce of our country no marked deviations are to be 
noted from the more common type. 

Anteriorly the manubrium juts out as a quadrate process with 
rounded angles; its inferior margin is continuous with a line that runs 
down between the slightly prominent carinal ridges, to become continu- 
ous below with the anterior carinal margin. 

Above, the general surface of the sternal body extends over it. A 
subcircular foramen, connecting the coracoidal grooves, pierces it at 
its base. The coracoidal furrow, thus becoming one groove, is biconvex, 
being depressed mesiad behind the manubrium, in which depression 
another pneumatic foramen usually occurs. Their upper and lower 



682 GEOLOGICAL SURVEY OF THE TERRITORIES. 

margins are produced slightly outwards, the inferior being the sharper 
of The two. 

The --eostal processes"' are exceedingly prominent, being bent over 
anteriorly at their apices, which are rounded. Behind them are observed 
the limited •■ costal borders.*' exhibiting the four transverse facets for 
the sternal ribs, and pneumatic foramina. The •• carina " or keel affords 
the greatest amount of osseous surface of the entire bone, greatly ex- 
ceeding the body. Its lower margin is a long convex curve outwards, 
and the ,k carina! angle" protrudes forwards nearly as far as the manu- 
brium, causing the anterior margin of the keel to be decidedly concave. 

The ■• carinal ridge" is thickened and heavy superiorly, where it 
limits or rather c nstitutes the boundary of the boue in that direction. 
Below it spreads out and is gradually lost, within the boundary of the 
carinal border proper. 

•• Subcostal" and " pectoral ridges" are nearly always well defined. 

The superior and interior xiphoidal processes are very characteristic 
of the Tetraon ida. They terminate by dilated extremities of nearly simi- 
lar shapes. Gupidania being an exception: the ends of the apophyses 
of the superior pair in this bird being rounded posteriorly (Plate XI, 
Fig. 82 5 see Plate XIII. Fig. 91. Lagopus. ifor the common pattern;) These 
processes arise from a common stem, and their shafts are flat internally, 
with a raised ridge extending the entire length externally. The •* body" 
of the sternum is. as a rule, very narrow, and notably concave anteriorly, 
becoming nearly flat behind, where it is produced beyond the keel for 
a greater or less distance. 

The manner in which this part terminates varies in the different 
Grouse. 

In Centrocercus if is nearly square across: in L a gop us roundly notched 
in the middle line, as it is in Bonasa : in Tetrao canadensis it is broadly 
cordate: while in Canace obscura. Gupidania, and Pedicecetes it is dis- 
tinctly cuneiform. The body is very narrow in Bonasa. approaching 
the Orti/gi/uc. where it seems really to be nothing more than a good 
ribbon-like finish to the superior border of the keel. In these birds, 
too. we are struck with the double carinal margins anteriorly formed by 
the projecting ridges, and the long spicula-form costal processes that 
extend nearly hall- way up the shafts of the coracoids. 

So much do the sterna of the Grouse resemble one another in species 
of average size that it would puzzle one not a little to tell them apart 
if they were separated from the skeleton, and we were not allowed to 
examine them in connection with other diagnostic features of the osse- 
ous parts of the species to which they might belong. 

In the Ortygince dilatations at the outer extremities of the xiphoidal 
prolongations are sometimes but moderately developed, as in our speci- 
men of Gyrtanyx rnassena. and the Plumed Partridge. The dilatations of 
the anterior or shorter pair of these processes are very broad mlleleagris. 
as are the stems that support them; the expanded part may have a 
foramen in it. or it may become bifurcated and the same specimen may 
show both varieties of termination. The longer or posterior pair in this 
species become very narrow behind, but stouter, and scarcely support 
any terminal expansions at all. The anterior half of the keel of the 
sternum of the Wild Turkey is very thick and strong, and all its ante- 
rior parts are prolonged upwards and forwards, being massive and lofty 
in very old birds. In other respects the sternum is stamped with the 
leading features of the bone as found among gallinaceous fowls generally, 
and this remark applies also to the sterna of Paro eristatus and Xumiela. 

Gupidania and Pedicecetes are particularly alike, but the former could 



snuFELDT.] OSTEOLOGY OF THE TETRAONHLE. 683 

be recognized by its superior xiphoidal processes, Bonasa by the nar- 
row body, Centrocercus by its size in the larger specimens, and so on. 

We will still continue to consider such of the vertebral column as is 
confluent in the old bird, or rather such vertebrae as become confluent 
and are more or less embraced by the ossa innominata, as the sacrum, 
and composed of sacral vertebrae, attempting to make no such divisions 
as Professor Huxley did, in his Anatomy of Vertebrated Animals, of 
this compound bone, though we must believe that this author is emi- 
nently correct in the view that he takes of this bone. 

There are sixteen of these segments that are to be so reckoned in 
Centrocercus, but it is only in the "bird of the year" that they can be 
counted with anything like accuracy, and even then great care must be 
exercised, and various pelves examined and compared with the younger 
birds at different stages and ages. 

The first sacral vertebra possesses free pleurapophyses, whose haema- 
pophyses do not reach the costal borders of the sternum, but articulate 
in a manner to be described further on. Regarding the pelvis from be- 
low in Centrocercus, we note that the anterior four sacral vertebrae have 
their combined par- and diapophysial processes thrown out as braces 
against the expanded anterior iliac wings. After this the ilia change 
their form to accommodate themselves to the basin of the pelvis, which 
they assist in inclosing, and with this change the succeeding vertebrae 
have their diapophyses much elevated to meet the internal iliac margins. 

This section consumes four additional vertebrae, the centra of which 
go to make up the latter moiety of the cavity for the u ventricular dila- 
tation" of the myelon, and they show the double foramina on either 
side, one above another, for the separate exit of the motor and sensory 
roots of the sacral plexus. 

A double row, i. e., one on either side of subcircular vacuities, exists 
here also among the transverse processes (Plate XIII, Fig. 90, Canace 
canadensis, $ ). It is through this portion of the sacrum that we observe 
in the chick the greatest amount of tardiness in sealing up of the neural 
tube above by the superior union of the engaged neurapophyses. 

The remaining eight vertebrae become much compressed with ex- 
panded processes that rarely allow apertures to remain among them, 
forming an excellent mid-section to the broad and capacious pelvic 
cavity, with nearly all signs of its original formation obliterated on the 
outer and superior aspect. 

The neural canal is distinctly circular as it enters the sacrum an- 
teriorly, becoming only slightly flattened as it nears the coccyx. 
Above we find the neural spine confluent with the ilia anteriorly along 
its summit, and some additional bone deposited posteriorly in the way 
of their lateral plates, to bridge over the ample "ilio-neural" canals. 
Opposite the "gluteal ridges" the bones are yet firmly knit, but for the 
remaining part of the sacro-iliac suture the interested bones can be 
said only to snugly meet each other. Cupidonia alone has quite an in- 
terspace present (Plate XII, Figs. 83 and 84). 

The sacral wedge is quite thoroughly permeated by air, which enters 
through foramina in the vertebrae in localities similarly situated to 
those described in speaking of the anterior part of the column. 

In Centrocercus sometimes the first sacral vertebra bears a well-de- 
veloped hypapophysis, and there may even be some evidence of this 
process on the second segment. The expanded extremities of these 
median processes are connected along their inferior margins by a deli- 
cate ligament of a firmer consistence than that membrane, which fills 
in the vacuities between the processes to which it is a limiting border. 



C84 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



£5ow, it depends how far this ligament is conducted backwards as to 
how many of the anterior sacral vertebras bear hypapophyses, as from 
its attachment to the hypapophysis (we have never seen it commence 
on the cervicals, though the directions assumed by their processes bear 
it out) of the first dorsal it completes a long shallow arc of an ellipse, 
in which the lower margins of the hypapophyses are found and assist 
to complete. This semi-osseous, semi-membranous, attenuated median 
plate dips down into the thoracic cavity in the living bird for some little 
distance, as an interpneumouic septum. 

The number of free caudal vertebras in the adult Sage Cock is five, 




Pedicecetes phasianellus. * 

and to these is to be added the pygostyle. They all have pretty much 
the same general appearance, differing principally in the length of their 
outstanding transverse processes. In this respect the first and second 
are about equal; in the third and fourth two or three millimetres are 
added on either side, with a roughened tuberosity above; while the fifth 
and last is in appearance and size very much like the first. These dia- 
pophyses are rounded at their extremities, and all slightly deflected. 
The centra are transversely elliptical and of good size. The neura- 
pophyses arise from them to inclose a subcircnlar neural canal, which is 
roofed over by the bifid and clubbed terminations of the elements, that 
hook or lean forward (Plate IX, Fig. OG; Plate XIII, Fig. 91). We 
have never observed an hypapoj>hsis upon any of them; and all the 
elements and processes, as always occurs in the cocygeal series, are 
more or less imperfectly developed. 

The pygostyle is an acute subcompressed triangle with tuberous base. 
Anteriorly and above it exhibts a bifurcated process that simulates the 
crests of the series ; below this a shallow facet, for articulation with the 
last caudal, and a feeble subcorneal depression to protect the termina- 
tion of the myelon between the two. Behind it has, at its lower angle, 
a deep groove, with a heavy bony rim or margin that shows a constric- 
tion near its middle (Plate IX, Fig. 65). This formation is easily ex- 
plained when we come to examine the development of this compound 
appendage in the chick of a day or so old. This will reveal the fact 



bhufeldt.] OSTEOLOGY OF THE TETRAONID^. 685 

that the pygostyle is composed of tliree vertebrae, which are fused to- 
gether, distorted and modified as the bird grows, so as to eventually 
result in the bone we have before us. 

We will now offer a few remarks upon the vertebral column, as ap- 
plied to others of the North American Tetraonidw. 

Of all the specimens examined, representing all the genera, the num- 
ber of distinctly free vertebrae between the skull and the anchylosed 
dorsals has been found to be invariably fifteen, the last one supporting 
movably-articulated pleurapophyses, that in Lagopiis, at least, possess 
small epi-pleural appendages. This portion of the spinal column, then, 
will constitute the cervical division, and in it we note that the third 
and fourth segments have interzygopophysial bars present, becoming 
so broad in the quails that the foramina they give rise to are sometimes 
very minute. 

The pleurapophyses become long and sharp in some and more con- 
spicuous than in others; this applies particularly to Cupidonia and Pe- 
dicecetes. The last cervical in the former has osseous spiculae leading 
backwards from its neural spine and postzygapopbyses, as in the anterior 
dorsals. In the same vertebrae the hypapophyses become confluent to 
form one broad process placed traversely with a mesial keel in front 
as the only indication of its original device. 

The normal number of cervical vertebrae in any of the species of Wild 
Turkey of this country is likewise fifteen, and this number holds good 
also, in the Guinea Hen and the Peacock. 

The carotid canal seems to commence and terminate in the same ver- 
tebrae both in Tetraonidw and Ortygince. 

Of the eminently characteristic dorsal bone little has to be said in 
addition to what has already been noted above. The number of verte- 
brae included in the anchylosis is never more nor less than four, and the 
first vertebral ribs have no haern apophyses; so they never connect with 
the sternum, though they invariably bear uncinate processes, and in 
many species contribute a good share toward the stability of the thoracic 
parietes. The metapophysial processes vary principally in their extent : 
in Bonasa, Cupidonia, and Pedioecetes. In several specimens they nearly 
cover the tops of the diapophysial arms. In Lagopus a narrow bar de- 
fines them, connecting the extremities of the transverse processes with 
comparatively few offshoots. In Canace canadensis the bone is very long 
and delicate. In C. obscura it has the same general appearance as in 
Centrocercus. The hypapophyses develop after the common type, but 
often irregularly ; they are found to be missing on the ultimate segment 
or rather the pelvic end of the bone in the Quails. 

The free and last dorsal may or may not have a hypapophysis ; we 
believe it never does in the Ortygince. It is overlapped by the ilia in 
Cupidonia and the Sharp-tailed Grouse. We find this to be a very 
marked feature in the last, and well marked among some of these birds 
sent me by Oapt. James (J. Merrill, Medical Department United States 
Army, from Fort Ouster, Mont., at a time they were particularly accept- 
able, and when this erratic fowl became suddenly and unusually rare in 
many localities, so that the gift was reckoned at the time as more than 
valuable desiderata. As far as the sacral vertebrae are concerned, the 
same general plan seems to be carried out : sixteen seems to be the al- 
lotted number among the Grouse and, as a rule, among the Partridges. 
In these birds the sacrum, i. e., the anchylosed sacral vertebrae, is much 
narrower, as is the entire pelvis, and much fewer perforations are to be 
seen among the dilated processes that go to meet the ossa innominata. 

In Citpidonia and Pedioscetes the sacrum is very broad, conforming to 



686 GEOLOGICAL SURVEY OF THE TERRITORIES. 

a pelvis in these birds that will be described further on. We find in a 
specimen of Canace canadensis, for which we are greatly indebted to Mr. 
Manly Hardy, of Brewer, Me., where the sacroiliac anchylosis is so per- 
fect, and original land- marks so obscure, that one might easily imagine 
the pelvis in this individual as being developed from a very much fewer 
number of ossific centres. The caudal vertebrae number five in all the 
Grouse except Cupidonia and Pedioecetes, these birds each having dis- 
tinctly six apiece. We would especially call the reader's attention to this 
fact, because when we come to discuss the pelves of these two birds, and 
recapitulate general skeletal data, it will be found that, as far as osteo- 
logical similarities are concerned, they come very near to each other. 
The coccygeal vertebra, otherwise, in common with the pygostyle, show 
very few differences worthy of record. 1 In Ortyx and Lophortyx there are 
but four caudal vertebrae and the pygostyle is markedly acute and long. 
Occasionally the last segment is but feebly developed, as in Lagcpus, 
where it may be a mere nodule ; and in Bonasa, too, sometimes a sixth 
vertebra can be be found, but usually requires force to detach it from 
the pygostyle, and in our specimens seems to be one of those that origi- 
nally formed that bone — though we do not deny in the face of such evi- 
dence that specimens of the Ruffed Grouse may be found that possess 
six of these vertebrae. 

Of the Scapular Arch and the pectoral limb.— Tins, arch, with its con- 
comitant, could have, with all propriety, been described in connection 
with its vertebra, but so distinct has it become, and so far removed in 
order to assist in carrying out such a notorious function as the bird's 
flight, that the author prefers to follow the general ruling of others by 
discussing it separately. Nothing could be more entertaining in the 
whole range of osteological research and study than the contemplation 
of the various avian shoulder girdles, with their attached wing bones, 
particularly the former, as exemplifying the law of equilibrium between 
a bird's habits, the never- varying part it is to play in nature, and its 
skeleton or the tramework that has been given it to carry that part out. 
This thought invariably enforces itself upon me in every instance after 
an examination of a collection of clavicles of different species of birds. 
It seems that there could not be an equipoise established anywhere in 
living 1 nature more thoroughly compensatory than that 'twixt a bird's 
power and mode of flight, and its scapular arch and other bones about 
the chest — to meet it, more essentially the clavicles. See the broad, ex- 
cessively pneumatic, yet robust, clavicular arch in any of the genus Ca- 
thartes, birds that sail aloft for hours apparently without fatigue, or the 
very similarly shaped arch in the Canada Goose, but in the latter for 
a very opposite style of sustained flight is non- pneumatic ; the feeble 
and often ununited arch in Speotyto, a bird with scarcely any preten- 
sions to being a good flyer at all ; in short one would, having a thorough 
knowledge of a bird's habits, be, in the vast majority of instances, able 
to guess very near as to the pattern of the furculum he would expect to 
find. 

Now we have seen, in reviewing the skeletons of the Grouse, that in 
many points some of the species, if we disregard size, similate each 
other very closely, as for instance in the various sterna and vertebrae. 
The clavicles of these birds form no exception to this rule, as far as 

1 The number of caudal vertebrae present or absent in any species must not be over- 
rated as a character, as the segments are liable to vary. I once heard it said that the 
number of true caudal vertebrae, including those that went to form the pygostyle, 
ought to equal the number of pairs of feathers in the bird's tail, judging from the ar- 
rangement as found in Areheopieryx. 



shufeldt.] OSTEOLOGY OF THE TETRAONIDtE. 687 

general appearance goes. The common model is seen in Plate XII, Fig. 
87, Cupidonia; but observe even here in these closely-related fowls how 
habit still tells upon skeletal characteristics. The broad, and not deep, 
pneumatic U -arch of Cathartes becomes the long non-pneumatic, almost 
acute, V-arch of the birds we are describing ; so, in view of being 
familiar with the habits of the Sharp-tailed Grouse and Sage Cock, 
need one be surprised to find in the fourchette of the first a depth of 4 
centimeters (omitting the inferior clavicular expansion), with an inter- 
space of 2.5 centimeters separating the superior articulating extremities, 
as compared with the bone in the second, where the depth is 6.2 centi- 
meters, and the interspace above only 1.7 centimetres. 

The bones of this arch are easily taken apart by maceration, and as 
has already been hinted they resemble each other very closely among 
the genera. The posterior angle between -scapula and coracoid averages 
for the species about 60°, and among the Grouse the distal end of the 
shoulder-blade is usually found to overhang the fourth dorsal pleura- 
pophysis on either side ; but in the California Quail these bones actually 
extend so far back as to overshadow the ilia for nearly a centimetre, 
they being proportionately narrow for their unusual length. This bone 
with the Grouse is only semi-pneumatic, its pneumaticity being confined 
to its coracoidal extremity, where the foramen is found in its usual site. 

The blade is thin behind, becoming stouter as it nears the glenoid 
cavity, sharp along its inner border, rounded without, clubbed at its 
pelvic end, and turned gently outwards along its entire length. Ante- 
riorly it contributes about one-third of the surface for the humeral joint, 
the coracoid the remaining two-thirds, while the acromial process within 
reaches forward barely to touch the clavicle, forming, as it does so, 
the usual interscapulo-coracoidal canal 

These bones are narrow in Bonasa and in C. canadensis, more curved in 
Lagopus, and very stout in Centrocercus, a good medium being seen in 
the Pinnated Grouse (Plate XII, Pig. 85). 

In Meleagris, they have their distal extremities cut square across, and 
reach back slightly beyond the pelvis. These bones in this species are 
also very stout and strong, and present the interesting variation in hav- 
ing a long pneumatic foramen on the superior aspect of each just beyond 
their glenoidal ends ; this feature is not present in Pavo. They are ex- 
ceedingly narrow and long in the Massena Partridge. 

The Coracoids are quite stout bones, but devoid of any marked 
peculiarities ; after leaving their sternal beds they extend upwards, 
forwards, and outwards, the furculum governing their distance apart 
at their upper ends. The glenoidal procss is extensive, and rises 
nearly as high as the head of the bone; the clavicular process is faintly 
bifurcated, and curls over slightly towards the fellow of the opposite 
side. The shafts are compressed from before backwards, and soon 
dilate into broad expansions below; anteriorly one longitudinal muscular 
ridge marks the surface of the bone, while on the sternal aspect two or 
three very distinct ones occur. The inner angle of the dilated base 
nearly meets its fellow of the opposite side through the foramen, at the 
root of the manubrial process ; from this corner outwards, for about 
two-thirds of the distance the surface is devoted to a curved articulating 
facet for the coracoidal groove, while the outer angle is tilted upwards 
and finished off by a distinctive little tubercle. On its posterior aspect, 
about the middle of the dilated part, a large, irregular pneumatic fora- 
men is found in all the Grouse, but seems to be absent in the Partridges. 
They are very large in Meleagris, at the same time being elliptical and 



688 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



so open as to give a pretty fair view of the interior of the bone in some 
cases. 

The position of the united clavicles, or the free acromial extensions of 
the scapulas, is shown for Lagopus in my drawing in Fig. 91. Something 
has been said about this bone already; we will add, however, that the 
superior ends always terminate by rather tuberous enlargements, smooth 
internally, but even as applied to the clavicular processes of the 
coracoids. The shafts are gently curved, of even calibre, and fall nearly 
directly downwards in some species, to be slightly expanded beneath in 
order to give better support for the large median dilatation below; this 
is triangular in outline, thickened in front, sharpened behind. In Ortyx 
this process is sometimes produced backwards, so as to nearly touch the 
sternum; this feature obtains, also, among some of the other Quails. 

The furculum of Meleagris is very different from that bonr, as just 
described in general terms for the Grouse. It is V-shaped, to be sure, 
but for the size of the bird is extremely slender, the medium plate is 
smaller than we find it in many of the Quails; the superior halves of 
the limbs are dilated and exhibit excavations on their inner aspects, at 
the bases of which we find irregular groups of pneumatic foramen; 
these limbs terminated above in truncate and thickened ends for artic- 
ulation with the remaining bones of the shoulder girdle. 

The furcula of Pavo and Numida are much more like the general type 
of the bone as found in the Tetraonidce than the same bone in the Wild 
Turkey is. 
The bones of the shoulder girdle are all well advanced in ossification 

in the young chick, 
but do not develop 
their distinctive 
markings until a bird 
is pretty well along 
in age ; this applies 
^\ more particularly to 
muscularlinesonthe 
shafts, the base of 
the coracoids, and 
the clubbed extrem- 
ities of the scapulae. 
In Bonasa, where we 
noticed how the body 
of the sternum was 
narrow like the 
Quails, we find also 
the median process 
of the united clavi- 
cles produced back- 
wards towards that 
bone. This Grouse's 
skeleton, in fact, 
seems to have the 
greatest tendency Partridge- ward over any other of the North American 
Tetraonidce. 

The free ossicle of the shoulder-joint, the os humero scapulare, is not 
present in any of these birds ; a firm piece of inelastic cartilage seems 
to supersede it and fulfill a like function. 

The humerus (Plate VII, Fig. 57, H— Figs. 60, 61 ; also Plate X, Figs. 




Lagopus albus. 



shufkldt.] OSTEOLOGY OF THE TETRAONID^. G31) 

7G, 77 of Cupidonia) of Gentrocercus is so exceedingly regular that it 
could be well chosen as the type of that bone in all birds in which it is 
pneumatic. It is due proportion for the size of the bird to which it 
belongs, possessing the usual sigmoidal curves from lateral and superior 
aspects (we describe the bone in situ in the closed wing) in graceful, though 
not decided prominence. The head or proximal extremity, slightly bent 
anconad, displays the most usual points for examination about it ; an 
extensive convex, smooth surface is seen for the glenoidal cavity, below 
it a deep notch, then the well-defined " ulnar crest" or lesser tuberosity 
curling over a large sub-elliptical pneumatic foramen, that is so patu- 
lous that the osseous trabecules and net- work are plainly seen at its base. 
The radial crest encroaches but very little upon the shaft, is quite stout, 
and only at its summit shows any disposition to curve over palmad. 

The shaft is smooth, elliptical on section throughout its extent, and 
almost entirely devoid of any muscular markings or lines ; it swells 
gently in the vertical plane as it approaches the distal end of the bone, 
upon which is placed the ordinary tubercles for articulation with radius 
and ulna. The inferior condyle, the internal of human anatomy, is the 
better developed of the two, and the ulnar tubercle is produced a little 
beyond the bone distally, while the oblique tubercle is brought down on 
the shaft. The olecranon fossa is rather shallow in the Grouse, being 
decidedly better shown among the Quails. There are no prominent 
points of difference in this bone among the Tetraonidce, except in regard 
to size ; Cupidonia, which has rather a heavy skeleton any way, the 
humerus is moderately robust in accordance. 

In the Ortygince the same characteristics are to be seen ; but we discover 
in addition at the proximal extremity of the bone, on its anconal aspect 
between the greater and lesser tuberosities, a deep fossa that has a great 
resemblance to the pneumatic orifice externally, and of about the same 
dimensions, being only separated from that depression by a thin bony 
wall ; it seems to be designed simply for muscular insertion, and has no 
communication with the general cavity of the hollow humeral shaft. 

The radius and ulna are also singularly typical in their avian char- 
acteristics, as might have been looked for after our remarks upon the 
bone of the brachium; their principal difference lies in their being non- 
pneumatic, although they are hollow like all long bones. 

In the right arm of Lagopus Uucurus, which we have before us, care- 
fully dried, in its position of rest, with all the ligaments still attached, 
in situ, we find the radius to be unusually straight, in fact almost inline 
between the oblique tubercle of the humerus and the bone it meets in 
the carpus. Its shaft is nearly cylindrical, and shows a muscular line, 
upon an otherwise smooth surface, that travels along its proximal two- 
thirds beneath. Distally it overlaps the cubit by a transversely dilated 
extremity to articulate with its carpal bone. The head of the ulna is 
large, and betrays the fact that it belongs to a bird of considerable 
power of flight ; the olecranon process is a blunt, tuberous apophysis, 
slightly bent anconad; the greater and lesser sigmoidal cavities are dis- 
tinct and fairly marked, particularly the former. 

The shaft of the bone is more than twice the bulk of that of its com- 
panion, decidedly convex outwards, the curve being greatest at the 
junction of the proximal and middle thirds; it is elliptical on section, 
the major axis of the ellipse being vertical. The muscular lines of the 
ulna are but faintly developed, as are the row of minute tubercles for 
the bases of the quills of the secondaries. 

Anteriorly the bone displays its usual trochlea head for its own carpal 
segment 5 this surface is bounded palmad by a sharp and even curve, 



690 GEOLOGICAL SURVEY OF THE TERRITORIES. 

convex distally. while the inner articulating surface beneath the ex- 
panded end of the radius is uneven and applied to the concavities and 
convexities of the free pair of carpal elements. 

These boues among the Partridges and other Grouse, except in size, 
present to us no extraordinary departures from the description just 
given of Lagopus. This remark applies also to the Wild Turkey, but the 
wing-bones in this bird seem to be shorter in comparison with its gen- 
eral size. 

We do not believe there is a bird in our country that can offer us better 
facilities for the study of the bones of the carpus than the young of 
Centrocercus uropliasian us. Anchylosis of the various segments involved 
is exceedingly tardy, and it is not at all necessary for the student of 
this joint, that has puzzled so many comparative anatomists and orni- 
thotomists, to seek the primoidal ossicles in the very young chick, unless 
he desires to ascertain the points as regards priority of ossification of 
the carpal bones, a question we will evade here entirely, for these bones 
are quite distinct and easily detached in the bird at six weeks or more 
of age. such as we offer our reader in the plates. 

In the adult Sage Cock, the carpus has the appearance of this joint, 
as it is seen in nearly all of the class where there are two free carpal 
bones, and the os magnum confluent with the proximal extremity of 
the metacarpus, and the mode of articulation is the same. This we 
know to be, first, a free, six-sided, uneven bone, the scaphoid, articulat- 
ing chiefly with the distal extremity of the radius and the metacarpus. 
This is the scaplio-lunar of my former papers, and we retain the same 
name for it here; it is also the rddiale of Prof. Edward S. Morse, who 
has made such positive advances in the elucidation of the tarsus and 
carpus in birds. 

The second bone is the cuneiform, larger than the first, and engaged 
principally by the cubit, but having also a process and an articulating 
surface for the confluent carpal and metacarpals ; this is the ulnare of 
Morse. 

These are the two carpals that remain free during life. 

We will now devote ourselves to the joint as observed in the bird at 
six weeks of age. We have no trouble in finding scaplio-lunar and cunei- 
form whatever. At the summit of the second metacarpal there is found 
a concavo-convex segment, that is universally taken to be the os mag- 
num, termed also carpale by Morse (Plate Til, Fig. 59. om). It articu- 
lates anteriorly with the upper end of the index metacarpal, covers the 
entire proximal extremity of the second, and nearly or quite meets 
another bone behind that is grasped by cuneiform ; this is the unciform 
(Plate Til, Pigs. 57 and 59. z). It has the appearance of being a detached 
and bulbous extension of the third metacarpal, and is about the size and 
shape of an ordinary grain of rice, having a shallow concavity on its 
anconal aspect. 

There is yet one perfectly free and distinct bone to be observed ; it is 
found on the inner aspect, very near the extremity of second metacar- 
pal, just below os magnum; it nearly meets unciform, and articulates 
with the process of cuneiform behind. This little segment is flat and 
very nearly circular, being applied by one of its surfaces against the shaft 
of the metacarpal, and held in position by ligaments. This segment we 
do not find described by any author known to us. and here call it the 
pentosteon. 1 The manner in which it eventually joins the metacarpus and 

1 This little bone of the carpus, which I still believe was originally described by 
myself, was called in the first edition of this Monograph the pisiform. This name 
might lead the student into the belief that I considered it either the analogue or even 



8HCFELDT.] OSTEOLOGY OF THE TETRAONID.E. 691 

the conformation it gives to that hone in the adult are shown in Plate 
VII, Fig. 58, and it is marked 8 in Fig. 59 of the young Sage Cock. 
The metacarpal for the index digit is likewise detached, although even 
at this early date it begins to assume a likeness to the bone as found in 
the adult. 

Medius and annularis metacarpals are also distinctly free, and their 
size and position faithfully portrayed in the figure alluded to above. 
The prominent process on the rear and upper third of the shaft of me- 
dius of the adult is now found only in cartilage in the younger indi- 
vidual. The rest of the bony part of the pinion is familiar to all of us ; 
it consists, in the adult "Cock of the Plains," as in all Grouse and the 
Ortygince, of a trihedral phalanx for the first metacarpal, constituting 
the index. We have found this phalanx in Meleagris and Ortyx bearing 
a well developed claw, covered with horn as in the ungual phalanges of 
of the feet; this is also the case in Numida and Pavo, and there is hardly 
a doubt but that this is the case with all of the American Gallince, and 
in this respect they agree with this class of birds of the Old World. Two 
more phalanges are devoted to the medius metacarpal, the superior joint 
or phalanx having the usual expanded blade, with the long free and 
pointed finger-bone below; and finally the smallest phalanx of all, freely 
attached to the distal end of the third metacarpal or annularis. 

These bones are non-pneumatic, as are all the bones beyond the bra- 
chinm in the Tetraonidce. 

Of the Pelvis and the Pelvic Limb. — After what we have said and seen 
in regard to the dilatory manner in which originally primitive elements 
in these birds anchylose, and only after the lapse of weeks condescend 
to amalgamate and form the confluent bones and cavities that occur in 
the major division of the Class, we must not be surprised to find the 
same routine and a like tardiness exhibited in the pelvic hsemal arch, or 
the pelvis, and its appendage the lower extremity. 

A glance at the figures illustrating the condition of the bones in ques- 
tion, of birds from one to two months old, will convince us at once that 
the rule still obtains ; in them we find the sutures among the ossa in- 
nominata still ununited, and the three bones of either side of the pelvis 
independent, and easily detached from each other about the cotyloid 
ring. 

In the young chick of Centrocercus (Plate VIII, Fig. 62), the ilium 
is a scale-like bone that rests against the sacral vertebrae. The pre- 
acetabular portion is excessively thin and delicate, and at this period 
constitutes the longest and widest part of the bone ; its border mesiad 
bears no vertebral impressions, but is sharp and brittle ; the outer margin 
is slightly rounded; this condition increasing as we near the diminutive 
acetabulum, where it is the stoutest. The distal margin, imperceptibly 
continuous with the inner, presents a convex curve anteriorly; the in- 
cluded surface viewed from above has a general concave aspect, espe- 
cially near its central portion. As we proceed backwards, however, it 
gradually becomes convex, to rise over the region opposite the cotyloid 
ring to form the general convex surface of the post-acetabular portion. 

The distal margin of the bone is nearly square across, and does not 
extend as far back as the ischium, the two being bound together at this 
age by cartilage, which has been removed in the plate. The outer and 
exterior margin of this division of the bone is rounded and fashioned 

the homologue of the pisiform as found in the Mammalia ; such at present being by no 
means the case, and all things considered, I have taken the liberty to change its name 
to the pentosteon, it being the fifth bonelet in the avian wrist. The same argument 
might be urged against the unciform, but I had nothing to do with the christening 
of that bone. 



692 GEOLOGICAL SURVEY OF THE TERRITORIES. 

to the cotyle, the anterior half of which it eventually forms. From this 
point it arches backwards over the fntnre ischiatic foramen. The inner 
margin of the post-acetabular portion is rather more rounded than its 
anterior extension, and directly opposite the ;; ring" presents for exami- 
nation the depressions of the tranverse processes of the ninth and 
tenth sacral vertebrae in elongated and concave facettes. The ninth 
vertebra seems to be the first to come in contact with the ilia, and the 
pelvis is so inclined that the remaining vertebra? soon, by their outgrow- 
ing apophyses, come in contact with it. ultimately fusing to develop the 
compound bone, usually termed sacrum. A good idea may be gained 
of the form of the ischium and os pubis by an inspection of Fig. 62. 
Plate VIII, at this stage of the bird's existence. For very obvious 
reasons the stoutest part of the ischiatic element surrounds the acetab- 
ulum ; an inferior process, extending towards a similar one directed 
backwards by the ilium, grasps the head of the os pubis between them. 
From this point the ischium sends backwards a triangular thin plate, 
rounded behind, that is intermediate in length between ilium and 
pubis. This latter is satisfied to follow suit and direction by contribut- 
ing a simple and diminutive spine. This crosses soon after the pubis 
has given its share to the cotylis. the obduratic notch of the ischium, 
converting it. as the bones fuse, into the obdurator foramen. 

TVe give an additional figure in the Plate, Fig. 63, that exhibits the 
further advances of these elements towards maturity. The bird is now 
nearly two months old. and it will be seen that one after another the 
vertebra? impinge upon the iliac margins within. The ilia and ischia 
extend behind, including vertebra after vertebra, from a series tha^ at 
this age might be easily taken for an extraordinary number of coccygeal 
segments. The head of the pubis at this time is entirely out of pro- 
portion with its rib-like extension, it having so spread and insinuated 
itself into the formation of the cotyloid ring as to form about one-fifth, of 
its circumference. At this stage, too, the necessary cartilage begins to 
be thrown out to form the future anti-trochanter on the ilium. 

So much for pelvic development ; now let us examine this bony basin 
as it is seen in the full-grown representatives of the genera. 

The manner in which the confluent sacral vertebrae meet the ossa in- 
nominata forming the ilio-neural canals and sacral sutures has already 
been defined above, so that here nothing more need be said of the mid- 
section of the pelvis taken as a whole. We give among the plates fig- 
ures of the pelves of Centrocercus. Canace canadensis: Lagopus. and Cu- 
pidonia from the skeletons of the adult birds, and if the reader will 
compare the lateral views given of the Ptarmigan and the Sage Cock 
he will see how few the differences are that exist between the two birds, 
and the same may be said, we assure you, for its superior and inferior 
views ; in fact in the case of Logapus it may be stated that its pelvis in 
most instances- is the perfect miniature of this bone found in Centrocer- 
cv.s in all respects. This applies, also, though not quite as strictly, to 
Canace and Bonasa. Of course certain minor differences are easily to 
be picked out, such as a greater fullness there, as slightly deeper depres- 
sion here, and so on: still our plate of Tetrao canadensis represents the 
general pattern of the pelvis among all the Xorth American Grouse, save 
Pedkecetes and CvpkJonia. In the superior aspect of this confluent bone 
in a fine specimen of Canace obscura, we find the pre-acetabular por- 
tions of the ilia very much depressed below the general surface of the 
bone, and quite concave. The convex anterior borders are embellished 
witlr a flattened rim that bounds them, often produced forwards as two 
median sharp-pointed processes. The post-acetabular surface is raised, 



SHUFELDT.] 



OSTEOLOGY OF THE TETRAONID.E. 



693 



and including the sacrum forms a gently convex surface ; the " gluteal 
ridges" dividing these two regions commence with the anterior iliac 
borders and are conveyed clear round to form the posterior ones of the 
same bones, describing two great Ss, the lower and outer curves of 
which pass by the acetabular projections, points where in some birds they 
terminate. 

Among the principal features to be noted in a lateral view is that 
the shaft of the pubis is in nearly all instances free from the ischium 
after quitting the cotylis and forming the elliptical obdurator foramen. 
If it anchyloses with this bone at all it usually occurs just behind that 
orifice. 

The ischium overlaps the pubis at two points — one quite broadly near 
its middle, and, again, by a process at its outer and inferior angle. 

The " ischial ic foramen" is the largest vacuity of the group of three 
that here present themselves; its boundary is sub-elliptical, with its 
major axis depressed posteriorly if it were produced. 

Both the internal and external margins of the cotyloid cavity, or ring 



rather, are circular, the former of which 



is not a little smaller, thus 



affording a very good and quite extensive surface for the head of the 
femur: the anti-trochanterian process or facet directed backwards is 
likewise ample, so that the femora are well supx^lied with articulating 
surfaces. 

There seems to be among the Grouse a predisposition for the ilia to 
overhang the region of the ischiadic foramina; it is most successfully 
carried out in Cupidonia. Viewed from below, we are struck with the 
amount of room and space these combined bones inclose; the profundity 
of the pelvic basin. This is very much enhanced by broad reduplica- 
tures of the ilia and ischia 
behind, and a general 
though even constriction of 
a prominent rounded border 
or rather ridge that extends 
from the fourth diapophysial 
abutment of the vertebra 
against the ilia on either side 
to the outer angles of the % 
ischia. Withi n, too, we often 
find about and at the base 
of these iliac fossae apertures 
for the entrance of air into 
these bones; such pneumatic 
foramina are also seen be- 
yond the os x>ubis and below 
the cotylis on either aspect. 

The pubic extremities 
never meet behind, though 
in many species they are 
very long and usually take 
the curve given them by the 
ischia just before leaving 
these bones. Their distal 
extremities are flattened in 
Ccntrocercus and generally more expanded than among the other varie- 
ties. 

The pelvis of Cupidonia is so different from the general description 
we have just given, that the author feels justified in giving his reader 




Ortyx virginiana. 



694 GEOLOGICAL SURVEY OF THE TERRITORIES. 

two additional figures, that present superior and lateral views of the 
bone in this bird; in comparing it with other figures given one cannot 
avoid being struck with these marked departures from the common type. 
There is one other Grouse that affects this style of pelvis, and that is 
Pedioecetes, and the attempt is not a bad one. The principal points 
wherein the Sharp-tailed Grouse has failed to make a perfect imitation 
of the unique pelvis of his ideal are, the ilia have failed to produce such 
ponderous overhanging lateral flaps, that nearly shut out from view the 
ischiadic foramen on either side. Again, these bones in Pediwcetes meet 
the sacral vertebra? for their entire length internally; and in this bird, 
too, the pubic elements often unite all along the inferior borders of the 
ischia. This does not occur in the Pinnated Grouse ; otherwise the 
bones are very similar and marked exceptions to the general pattern of 
the other members of this subfamily. 

After examining a large number of the pelves of our Grouse, and 
noting" their capacious cavities and great width from side to side, one 
cannot help but be surprised at the complete change in outline, on turn- 
ing to these bones as they are found among the Partridges. 

As a rule in these birds, the pelvis is elongated and unusually narrow, 
though the drooping ischia behind give it additional depth in its more 
posterior parts ; the pubic bones are turned up behind after they extend 
beyond the elements above them, and the lateral walls formed by the 
ilia are, just beyond the ace-tabula, almost vertical ; in fact, one would 
almost suspect the pelvis of the common Virginia Partridge as belong- 
ing to the skeleton of some variety of Curlew, had he not been sure of 
the owner. This apparent departure from the more general model of 
this bone in gallinaceous fowls, however, is not nearly so decided in 
other varieties of the Ortygince, as for instance in Lophortyx californica, 
and a very good drawing of the pelvis of this bird can be seen and 
studied, in Mr. T. C. Ey ton's Osteologia Avium, London, 1867, Plate 
22, figs. 1, 1, 1, 1 ; three of the figures here cited, however, are for the 
palatine bones, inferior view of sternum, and anterior and posterior 
views of the tarso-metatarsus. We have carefully compared the diam- 
eters of these representations with the skeletons of the species in ques- 
tion, and find them quite accurate. 

The pelvis in the Wild Turkey has the general resemblance of this 
bone as found among the gallinaceous birds at large. The sides of the 
ilia, in the preace tabular portion are nearly vertical, spreading out only 
anteriorly into the horizontal plane. We are struck with the unusual 
size of the ilio-neural canals. These passages are over three centimeters 
long, having a complete bony septum dividing them, composed as usual 
of the neural spine of the leading sacral vertebra?. The major part of 
the common roof above is formed by the meeting of the ilia. The post- 
acetabular region is quadrilateral in outline, rather extensive and very 
flat. In this locality the iliac borders meet the vertebra?, but only par- 
tially to anchylose with them. The sides of the pelvis are broad and deep, 
and the pubic bones never join with their lower margins. Just beyond 
the cotyloid ring, on either side, we find a stumpy spine, very much the 
same thing as we see in the common fowl. Between the ischiatic and 
obdurator foramen a deep triangular pit occurs, that usually has a few 
pneumatic foramina at its base. Five caudal vertebra? are found in 
the Wild Turkey, and the pygostyle is very long and sharp, being pro- 
duced backwards. In the Peacock we find the pelvis shorter than in 
Meleagris and generally arched above, as if the bone in this locality had 
gradually been distorted by the burden of feathers it bears above which 
pull upon the muscles overlying it. To afford additional support for 



6HUFFXDT.] 



OSTEOLOGY OF THE TETRAONID.E. 



695 



this beautiful appendage in this fowl, we find the four coccygeal verte- 
brae very broad and spreading, while the pygostyle is equally exten- 
sive, being composed of two horizontal plates connected by a vertical 
one, lying longitudinally in the middle plane. The pelvis and the caudal 
vertebrae, including the pygostyle in the Guinea Hen, are very much the 
same pattern as we find them among the Grouse generally, and both this 
bird and Pavo have the characteristic spine protruding anteriorly beyond 
the acetabulum on either side. 

The femur in the young chick of Centrocercus is but partially devel- 
oped; above, the head is almost entirely cartilage, while below the con- 
dyles are very indistinct and the bone bears no signs of pneumaticity. 
In a few weeks, however, these points rapidly exhibit tnemselves: a 
rounded trochanterian ridge is thrown out; the head essays to assume 
its sphericity; the condyles become evident ; the fibular groove appear- 
ing last of all and about the same time with the vascular foramen or 
medullary orifice at junction of upper and middle thirds. In our exam- 
ination of this bone in a fine old cock Cupidonia, and comparing it with 
others, we find that it is remarkably well balanced in point of length 
and general development. 

The trochanterian ridge is prominent and arches over the articular 
facet for the ilium; the neck is distinct and makes an angle of 45° with 
the shaft; the head is well formed, spherical, and in all Grouse seems 
to bear a double depression for the ligamentum teres. Anteriorly below 
the trochanterian eminence there is an extensive collection of pneumatic 
foramina. The " trochanter minor" never develops. 

The shaft is smooth, bent slightly forwards ; displays the usual mus- 
cular lines and the medullary orifice; it is nearly cylindrical on section 
about its middle, and before terminal expansion takes place. Below, 
the rotular channel is deeply grooved, separating the prominent con- 
dyles ; of these the external and lower one presents the usual fibular 
fissure ; behind, the popliteal depression is well sunken, one of the mus- 
cular lines running into it, and often a foramen is found at its base. 
Shallow fossae are found laterally at the outer surfaces of the condylar 
enlargements, and some- 
times a notch where shaft 
meets the internal one ante- 
riorly. The bone is usually 
slenderer in Bonasa, Pedi- 
cecetes, and Canace canaden- 
sis. 

Among the Ortygince the 
femur has the same general 
characteristics; it is, how- 
ever, non-pneumatic, the 
double pit for the ligamen- 
tum teres is better marked, 
and the muscular lines are 
scarcely perceptible. 

The patella is never absent 
in the Tetraonidce, and occu- 
pies itS USUal position as a Orortyx pieta. 

free bonelet protecting the anterior aspect of the knee-joint. It accom- 
modates itself to the conformation of the rotular channel, having a flat- 
tened surface superiorly, a rounded. border below, and a double surface 
behind, the most extensive aspect of which is applied to the side towards 
the internal condyle. 




696 GEOLOGICAL SURVEY OF THE TERRITORIES. 

The proximal extremity of tibia in the young chick of Centrocercushas 
advanced so little towards assuming any of the definite characteristics 
of the full-grown bird, that, almost in self defense, we take up for exam- 
ination the bone from a skeleton of a bird of the same species several 
weeks older : here we discover the superior general condylar surface 
still capped with cartilage, and the borders confining it, as yet, but 
feebly produced. The most interesting point, by far, is the appearance 
of an unusually large epiphysis, if it may be so termed, fashioned to 
and resting upon the future location of the "rotular crest." 

Why this bone should be here added we cannot, as far as our knowl- 
edge extends, exactly comprehend, for in the old and mature birds of 
any of the Grouse the epi-cnemial crest is never very prominently pro- 
duced, nor is it in any of their near kin. As age advances this seg- 
ment becomes thoroughly confluent with the tibia, and leaves no trace 
of its early existence. 

The head of the bone in the adult Sage Cock is a very substantial 
affair, with pro- and ecto cnemial ridges, well produced, that soon merge 
into the shaft ; the latter ridge is usually dilated on its anterior as- 
pect, and the rather extensive concavity between them is directly con- 
tinuous with the shaft below. 

The tibia never becomes pneumatic either in the Grouse or Quails, 
and in the former, sections of its ishaft are universally transversely 
oval ; the fibular ridge of the upper and outer third of this portion of 
the bone in Centrocercus is about 2 centimetres long, and appears to be 
little more than a raised and roughened line. Below the fibular ridge 
we find the nutrient foramen, but otherwise the tibial shaft is very 
straight and almost entirely devoid of any markings, at least to that point 
anteriorly, where the ascending groove coming from between the con- 
dyles impresses it, and that, with an increasing intensity to its termi- 
nation over the intercondyloid notch ; at this point a bony bridge is 
thrown across obliquely, the outer abutment of which is the lower. (Plate 
IX, Fig. 69.) 

The most engaging points of interest, so far as the tibia is concerned, 
center about the distal extremity of the bone. After careful examina- 
tions of the recently-killed subjects, dried skeletons, and carmine-stained 
specimens, the following result seemed to be presented with greater or 
less distinctness in every case, and these results correspond very nearly 
with Professor Morse's invaluable investigations and studies: In the 
young of Cmtrocereus, several days after leaving the nest, we observe 
at the future site of the tibial condyles, encased in the then articular 
cartilage, on either side, a free reniform ossicle. These ossify to the 
surface in time, and the outer becomes the fibular e, the inner the tibiale; 
both are tarsal bones. It will take time and further research to decide 
definitely as to which is the os calcis, and which the astragalus. Above 
the former, and a little towards a mid-shaft position, there is yet another 
free ossicle ; it is the intermedium of Morse. That it is another tarsal 
bone, there can be no doubt, we believe, at this date; but before we 
decide upon its special homology we must satisfy ourselves by further 
research and dissection of the young at various stages of development 
in birds and other vertebrates where this bone constitutes a feature of 
the skeleton. 

It soon fuses with the tibiale, and the latter with the Jibulare, so that 
the faintest traces are alone discernible in the bird at two months of 
age. (Plate IX, Fig. 67, E'). At this period the subsequent bony span 
for tendinous confinement above them has not commenced to ossify. 

In the adult the apex of the intermedium affords attachment to the 



6HUFELDT.] 



OSTEOLOGY OF THE TETRAONID.E. 



697 







oblique ligament that is attached higher up on the inner aspect of the 
shaft, that also holds some of the extensor tendons in position. The 
condyles in mature birds have an anteroposterior position at the 
extremity of the tibia ; these are of a uniform outline, and the inter-con- 
dyloid notch that separates them is of no particular depth until it arrives 
on the anterior aspect of the bone. Externally and laterally almost 
within the limits of the outline of the outer condyle we find two tuber- 
cles, one above another; the lower is for ligamentous attachment, the 
upper is the remnant and only existing evidence of the lower extremity 
of the fibula. A similar tubercle is found on the opposite, side corre- 
sponding to the lower one just described on the outer aspect. 

The fibula is freely de- 
tached and never completely 
anchyloses with the tibia. 
Its proximal extremity is 
clubbed, enlarging very 
much as it rises above the 
condylar surface of its com- 
panion from the fibular ridge ; 
i t is laterally compressed and ^^ 
convex above at the summit. 
Tn many Grouse the attenu- 
ated remains of its extension 
below can be traced on the 
shaft of the tibia, which bone 
has nearly absorbed this 
third of its weaker associate. Caiupepia squamata. 

In the early life of the chick of the Grouse we have been discussing, 
the combined tarsals are surmounted by a third plate of cartilage, that 
subsequently ossifies, apparently by one center. The bone thus formed, 
the centrale, we believe undoubtedly to represent either a single tarsal 
element or the connate bon s of the second row. 

At this age the metatarsals that combine to form the shaft of the 
tarso-metatarsus are still easily individualized, though well on the road 
toward permanent fusion. It will be observed that we still retain the 
term tarso-metatarsus. and we think justly so, as the compound bone of 
the mature bird has combined with it at least one of the tarsal bones. 
The tibia could with equal reason be termed the tibio-tarsus, and again 
the compound bone in manus, the carpo-metacarpus, but for obvious 
reasons such innovations are not always advisable. 

We discover in Cent rocer ens and Canaee canadensis — in that strong 
inelastic cartilage that is found at the back of the tarsal joint in all the 
Grouse, on the inner side — a concavo-convex free bone, nearly a centi- 
meter long in the Sage Cock, and two or three millimeters wide; this 
ossicle must be regarded only as a sesamoid, though it is nearly as large 
as the patella, and in no way as constituting one of the tarsal bones. 

It will be remembered that in the first edition of my monographs 
upon the osteology of Speotyto and Uremophida, the old term of the " cal- 
caneal" process was retained for that prominent projection found at the 
superior and hinder end of the bone tarso-metatarsus. It having any- 
thing to do with the calcaneum or the os calcis in the homologies of the 
avian tarsus, was stoutly denounced in the first appearance oi' my mon- 
ograph upon the osteology of the North American Tetraonidce. In these 
papers, as well as in my osteology of Lanius ludovicianus excubitorides, it 
was given the name of the tendinous process, from the well-known fact 
that bv one means or another it transmitted the flexor tendons at the 



698 



GEOLOGICAL SUEVEY OF THE TERRITORIES. 



be the case among all birds? 




back of the tarsus. At the present writing we know of no author who 
has very recently added anything to the literature of this subject, bear- 
ing upon this particular point, and the writer was quite confident that 
all he had said in his Grouse paper could be sustained ; for if this process 
did not separately ossify in the Tetraonidce, and thus weaken its claim 
as being a homolonge of one of the tarsal bones, why should this not 

A few months ago we made a discovery 
that again shook our faith in this 
matter, and led us to believe that 
perhaps this process was more 
entitled to the appellation of the 
calcaneal than Ave had previously 
supposed. The facts in the case 
are these, and we must leave our 
reader to draw his own conclu- 
sions. We have before us the 
tarso-metatarsus taken from the 
young of two widely-separated 
genera and families. The first 
is that of the chick of Pedicecetes, 
one of our present genera; the 
second is from the young of Gin- 
clus mexicanus, the American 
Water Ousel. In the first we 
observe all those characteristics 
Lophortyx calif omica. that we have already ascribed to 

this family in general, and elucidated above ; but in the tarsal joint of the 
Dipper, our second example, we find that the centrale and the so-called cal- 
caneal process are connately developed; the latter element being of good 
size already shows the grooves for the flexor tendons. Of the number of 
points of ossification in this development in Ginclns the author can say but 
little, as his material has been chosen from a few of these birds, furnished 
by the Smithsonian Institution. The subject is an extremely interesting 
and important one, and will well repay further investigation and research, 
and the author only regrets that at this date he can make but such 
slight additions to what we already know of it through the exertions and 
examinations of others. In any of our adult Grouse, this process is verti- 
cally grooved and perforated for the passage of tendons ; from its inner and 
posterior angle in many of the Tetraonidce it sends down a thin plate 
of bone that usually meets the shaft at junction of upper and middle 
thirds, occasionally running further down to become confluent with it in 
every instance; this feature is rarely present in the Quails. In Centro- 
cercus and others the hinder aspect of the tarso-metatarsus is sharply 
marked by muscular ridges. The superior and articulating surface of 
this bone displays eminencies and depressions fashioned to accommodate 
themselves to the condyles of the tibia ; a tuberosity on the anterior mar- 
gin in the articulated skeleton fits into the intercondyloid notch of the 
bone. Below this the shaft in front is scooped out, having at the base 
of the depression two small elliptical foramina, side by side, and to the 
inner side of its boundaries one or two pointed muscular tubercles. The 
distal and transversely expanded end of the bone presents the foramen 
for the anterior tibial artery, occupying its usual place, and the three 
trochlear apophyses for the toes, the middle one being the largest and on 
the lowest level. The two lateral ones, separated by wide notches from 
the former, are thrown but a limited distance to the rear, so that the 
concavity behind them is not peculiar for any great amount of depth. 



shufeldt.] OSTEOLOGY OF THE TETRAONIILE. 699 

The os metatarsale accessorium is situated rather high upon the shaft, 
and bears more than an ordinary semblance to a demi phalanx, with its 
distal articular trochlea. As usual it is freely attached by ligaments. 

The internodes are based upon the more common plan as applied to the 
avian foot; i. e., in the order of the phalanges, from the first to the 
fourth, 2, 3, 4, 5 joints. They are in proportionate sizes for their sev- 
eral owners, and possess nothing peculiar about them, having all the 
usual characters as found in the gallinaceous foot generally. 

They possess the usual enlarged and biconcave proximal extremities, 
with the distal and convex bitrochlear ends, with a more or less subcylin- 
drical shaft ; the ungual joints being but moderately curved doAvn wards. 

There are but few or no striking differences to be noted as existing 
among the lower extremities of our Tetraonidce. 

The bones are very delicately fashioned in Canace canadensis and the 
Sharp-tailed Grouse; that is, the calibres of their shafts seem to be less 
as compared with their general lengths, but they belong, we must remem- 
ber, to very trim little game birds, as contrasted with our heavy and 
ponderous old Sage Grouse of the western prairies. Our specimens of 
Lagopus and Canace obscura do not show the bony extension from the 
tendinous process at the back of the tar so- metatarsus , apparently present 
in all the others and alluded to above — Centrocercus sometimes proving 
an exception — and this bone never normalty develops a spur in any of 
our North American Grouse, as seen in birds of near kin. 

Tendons of the anti-brachium and pinion are very prone to ossify, and 
one is quite constant on the anterior aspect of the metacarpus. This 
applies with still greater force to the lower limb, where it seems that 
every tendinous extension of the muscles of thigh and leg become bone 
for their entire lengths, then forking sometimes over the fore part of 
tarso-metatarsus as they branch to be distributed to the podium. 

It may be found that Bonasa can claim normally six segments as rep- 
resenting the caudal vertebrae, and we have in our possession a pelvis of 
this bird where a rudimentary second sacral rib is evident, but this can 
only be regarded in the lighfi of an anomaly. 

The pelvic limb of the Wild Turkey has nothing about it of peculiar 
interest, i. e., in the adult fowl, and we have never been so fortunate as 
to examine the young of this bird. The entire extremity is non-pneu- 
matic, and composed of quite powerful bones. From the lower and 
outer angles of the tendinous process of the tarso-metatarsus, a thin 
plate of bone is thrown down that gradually approaches but does not 
touch the shaft until it arrives opposite its middle, when it merges into it. 
The lower portion of this lamina bears the short spur in Pavo, but of 
my three specimens of Meleagris, no such appendage is seen ; they are 
no doubt all females or perhaps young males before the development of 
the spur, as we believe old male Turkeys, always possess it. 

Interesting osseous malformations are occasionally to be seen, but they 
are beyond the jurisdiction of this monograph to treat; nor will it be 
practicable to enter into the engaging subject of the differences between 
the pelves in the male and female birds, but that such differences do 
exist there can be no reasonable doubt. 

Our Grouse and Quails all belong to the Alectoromorphce of Profes- 
sor Huxley's classification, sharing in common with the representatives 
of that group many of their osteological characters; some of these 
characters, such as the general form of sternum, certain cranial peculi- 
arities and others, are positively distinctive and differ from all other 
groups. Our Tetraonidce, with the exception of Pedicecetes, and Cupi- 
donia, have certain similarities or osteological resemblances found 



700 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



among the more prominent bones of the skeleton, as the pelvis, ster- 
num, and scapular apparatus, that bind the remaining genera very 
closely together. The same may be said in a general way of the Quails, 

or Ortyginw, taken as a subfamily, have 
many such characters in common, as 
compared among themselves. 

Cupid onia and Pedicecetes although 
they possess the usual tetraonial oste- 
ological characters referred to above, 
have in addition quite a number of mu- 
tual external characters, that compel 
us to regard these two forms as being 
of very near kin, and our osteological 
studies of the sub-family have certainly 
demonstrated the fact that this relation- 
ship is by no means weakened by the 
latter investigation. In short, although 
ornithologists will no doubt always retain these two forms in separate 
genera as the classification of birds goes, still it may be well to bear in 
mind that nearly or quite all of the anatomical characters of Cupidonla 
and Pedicecetes when compared together bring these two Grouse nearer 
to each other than any other two forms of the group in our fauna; so 
near, in fact, that but iittie violence would be perpetrated by restrict- 
ing them both to one and the same genus, and no doubt there are not 
a few instances in our present classification of birds where forms not so 
nearly related as these two Grouse are that have been retained in one 
genus. 

Washington, D. 0., May 1, 1882. 




Cyrtonyx massena 



702 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE V. 

Fig. 47. — Cranium of young Centrocercus. three or four days old, viewed from above. 

Fig. 48. — The same ; lateral view, with lower jaw added, showing articular element 
detached. 

Fig. 49. — The same, viewed from below. 

Fig. 50. — Centrocereus. Lateral view of skull of "bird of the year" four mouths old : 
the hyoid arch has been detached. 

Fig. 51. — The same aged bird as in Fig. 50. showing the disarticulated segments of 
the four cranial vertebra?: OV. neural arch of the occipital vertebra (epencephalie 
arch), first of the skull: OV. its lueuial arch in outline (scapular arch): so, super- 
occipital; eo, exoccipital ; po, the connate paroccipital: bo, basioccipital : FY. neural 
arch of the parietal vertebra (mesencephalic arch), second of the skull; P. V.'. its hietnal 
arch (the hyoidean): P. the parietal: ms, the mastoid: as, the alispheuoid; bs, the 
basi-sphenoid: gh, theglossohyal: cli, the ceratohyal ; blu the basi-hyal ; uh, theurohyal: 
hb and. cfe, the hypo-branchial and cerato-branchial elements of the thyro-hyals, 
respectively: F. v., neural arch of the frontal vertebra (proencephalic arch), third of 
the sknll : F. V.', its ha?mal arch (the mandibular): Fr. the frontal: x, the site of the 
postfrontal in some of the class: os, the orbito-sphenoid in ontliue; ps. the basi-pre- 
sphenoid; Tp, the tympanic; ar, the articular; 8. an. the surangular; an, the angular; 
se, the splenial element; and de, the dentary element: X. V.. the neural arch of the 
nasal vertebra (rhinencephalic arch), the fourth and last in the skull; X. V.', itsba?mal 
arch (the maxillary); N, the nasal; Pf, the prefrontal (ethmoid); F>, vomer in out- 
line, as it does not occur in this bird, (see description in text) ; PI, palatine ; mx, max- 
illary: i. »!jf, intermaxillary (or premaxillary) : 1. the petrosal: 2, thesclerotals; 3, the 
lacrymal; Pty, the pterygoid, the diverging appendage of PI, the palatine : sq and via, 
the squa7HOsal and malar, respectively, are diverging appendages of Tp, the tympanic. 



704 GEOLOGICAL SUEVET OF THE TEEEITOEIES. 



PLATE VI, 

Fig. 52. — Lateral view of skull of Cenirocercus, adult £ , hyoid arch removed. Tigs. 
54. 55. 58, 61. 64. 65. 66, 68, 69. and 70 are from the skeleton of the same specimen. 

Fig. 53. — Sternum of the chick, three or four days old: Cemtmeat 

Fig. "4. — Sternum of Centrocereus. viewed from below: adnlt $ . 

Fig. 55. — Dorsal vertebra?, lateral view, left side, from the same. 

Fig. 56. — Sternum of Centrocercus, '"bird of the year" (two months old), showing 
development of this bone: 4, !opko*tton; 5. metosteon; 6, pleurosieon. 



706 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE VII. 

Fig. 57. — Right pectoral limb of Centroeercus, same bird as the steinuin in Fig. 56 
was taken from. H, humerus, palmar aspect; r, radius; u, ulna; s, scaphoid (ra- 
diale) ; c, cuneiform (ulnare) ; z, unciform ; d, iudex digit ; 9, third metacarpal (an- 
nularis) ; 9', second metacarpal (medius); d', d", phalanges of the second metacar- 
pal ; d" ', phalanx of the third metacarpal. 

Fig. 58. — Left metacarpus of an adult Centroeercus. 

Fig. 59. — Left metacarpus of the same bird as figured in 57, showing all the seg- 
ments that go to make up the bone in Fig. 58 : om, carpale or os magnum ; z, unci- 
form ; 7, index or first metacarpal ; 8, pentosteon ; 9', second metacarpal ; 9, third meta- 
carpal (annularis). 

Fig. 60. — Left humerus (Centroeercus), taken from the same bird as in Fig. 57. 

Fig. 61. — Left humerus, anconal aspect, Centroeercus; adult. 



708 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE VIII. 

Fig. 62. — Ossa innoniinata, Centrocercus, three or four days old, showing how the 
pelvic hones form the acetabulum : X, ilium; Y, ischium ; Z, os pubis. 

Fig. 63. -^Pelvis, same bird as in Fig. LT; Centrocercus ; X, ilium; Y, ischium; Z, 
os pubis. 

Fig. 64. — The perfect pelvis, lateral view,, as in 62 and 63, of Centrocercus; adults 



710 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE IX. 

Fig. 65. — Posterior view of pygostyle, adult Centrocercus. 

Fig. 66. — The same, left lateral view. 

Fig. 67. — Pelvic limb, anterior view, Centrocercus, taken from same bin] as in Fig, 
57 : E, Epicnemial epiphysis of the tibia; E', the confluent tarsal bones found at the 
distal extremity of the tibia at this age. tibiale ( astral agus), and fibnlare (os calcis) : 
E", the centrale. 

Fig. 68. — Left tarso-metatarsus, inside view, adult $ , Centrocercus. 

Fig. 69. — Anterior view of right tibia and fibula, from the same skeleton as 68. 

Fig. 70. — Anterior view of right femur, same bird, as shown in Figs. 68, 69. 



712 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE X, 

Fig. 71. — Right lateral view of skull of adult $ Cupidoni'a cupir!<?. 

Fig. 72. — Lower mandible from the same, viewed from above. 

Fig. 73. — Skull from the same, lower jaw* removed: seen from above. 

Fig. 74. — The same from below. 

Fig. 75. — Sclerotals, right eye from the same. 

Fig. 76. — Right humerus, from the same, palmar aspects 

Fig. 77. — The same, aneonal aspeet. 



71-4 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XI. 

Fig. 78. — Sacral vertebral rib with its haernapophysis, left side : w'. the pleurapophy- 
sis. posterior view: ;r. the corresponding haernapophysis. 

Fig. 79. — Fifth pleurapophysis with its corresponding haernapophysis attached : 
from the same bird, dorsal vertebrae, inside view. 

Fig. 60. — Fourth pleurapophysis with its corresponding haernapophysis attached ; 
from the same bird. [Cmpidimia), dorsal vertebrae, outside view. 

Fig. 61. — Sternum from below ; same bird. 

Fig. 82. — Sternum, left lateral view; same bird (Cupidonia cupido). 



71G GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XII. 

Fig. 8>. — Pelvis from above ; Cupidonia cupido. 
Fig. 84. — Pelvis, right lateral aspect; same bird. 
Fig. 85. — Right scapula and coracoid, in situ; same bird. 

Fig. 86. — Left coracoid, posterior view, from the scapular arch of the same bird. 
Fig. 87. — Right lateral view of clavicles, from the scapular arch of the same bird; 
dotted lines show the outline viewing it from behind. 



71 8 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XIII. 

Fig. 88. — Right lateral view of skull of adult $ Lagopus leucurus; hyoid arch has 
been removed. 

Fig. 89. — Cranium of Pedicecetes phasianellus ; lacrymal, nasal, and intermaxillary still 
attached. 

Fig. 90. — Pelvis, adult $ Can ace canadensis, viewed from above. 

Fig. 91. — Portion of skeleton of Ptarmigan, Lagopus leucurus, showing thoracic and 
pelvic bones in situ, with the scapular arch and caudal vertebrae. The last cervical 
vertebra still remains attached in order to show its free pleurapophysis. 



OSTEOLOGY OF LANIUS LUDOVICIANUS EXCUBITORIDES. 



By E. W. Shufeldt, M. D., 

Captain, Medical Department, United States Army. 



Mr. Robert Ridgway, in his carefully prepared check-list of North 
American Birds (Bull. No. 21, Nomenclature of N. American Birds, 
chiefly contained in the U. S. Nat. Mus., Washington, D. C), gives us 
the .representatives of Laniidwj the species borealis and ludovicianus, 
with the two western varieties of the last, robustus and excubitorides, 
which latter form we have chosen as the subject of this paper to demon- 
strate the peculiarities of the skeleton of these interesting birds. The 
habits of the Shrikes are well known to all ornithologists, so one will not 
be surprised, after a view of Fig. 100, in the plate (where our subject has 
been made, by the aid of the dissecting knife and maceration, to exhibit 
one of the truest indices of his character), to find the large, semi-hawk 
like skull surmounting the remainder of a bony frame-work, that might 
easily be mistaken as belonging to a Thrush or any other Osciue; but 
it is this very characteristic that individualizes these truly passerine- 
raptorial birds. 

In the skull, divested of the lower jaw and hyoid arch in the adult, 
we find that anchylosis of the primoidal segments has been very thorough ; 
outside of the bony parts of the sense capsules — the ossa quad rata and 
the pterygoids are the sole escapers of this notorious feature in avian 
craniology — indeed, we discover in the skull of this species, before it 
has left the nest, that the primitive segments that go to make up the 
group of bones that we described as the occipital vertebra in former 
papers are well advanced towards permanent union, especially about the 
condyle, traces of its formation being extremely difficult of discernment, 
and in the mature bird this hemispherical facet for the atlas is exceedingly 
diminutive, measuring only .5 of a millimeter in diameter. About the 
basi-cranii we find the usual foramina for the exit and entrance of vessels 
and nerves, and note in our examination that the anterior apertures of 
the Eustachian tubes are double, very small, and protected by an osse- 
ous lip from the basi-sphenoid The foramen magnum is sub-circular 
and of medium size ; together with the basi-cranii, it makes an angle with 
the horizontal plane of 25°, the anterior bearing point being the tip of the 
beak, and the two posterior bearing points being the internal facettes 
upon the ossa quadrata. That part of the cranium above, formed by 
the frontals and parietals is very broad and smooth, and quite often 
the sutural traces are easily made out, and in cases where maceration 
is persisted in, the coronal suture may gape; beyond, the interorbital 
region becomes slightly depressed. The pseudo fronto-maxillary articu- 
lation is denoted by a transverse line nearly a centimeter long, and is 
moderately flexible; the superior tips of the lacrymals form its lateral 
boundaries. The superior mandible is made up of the usual bones, it 

719 



720 GEOLOGICAL SURVEY OF THE TERRITORIES. 

is very broad at its base, and gently deflected throughout; the nasal 
bones bound posteriorly on either side vacuities that lead into the rhinal 
chambers, but the true nostrils are found beyond these, as distinct 
elliptical apertures. It is, however, the horny integumental sheath that 
really gives to this bird's beak its peculiar raptorial aspect, for when 
stripped of this, the osseous toinia show no sign of tooth or notch be- 
yond. Below, the palatine fissure is quite wide, and through its open- 
ing we discover that the ethino- turbinate are more or less developed, 
together with a partial septum narium, and the space is further intruded 
upon by a sub-compressed and originally distinct vomer, that is bifur- 
cated behind to receive the rostrum of the basi-pre-sphenoid, lodging a 
portion of the prefrontal in its fissure above. 

The palatines have become amalgamated with the inter-maxillary 
anteriorly, and form, with the pterygoids, the usual joint on the rostrum 
of the sphenoid behind ; they throw out sharp lateral apophyses that 
are directed backwards. The pterygoids are very much expanded at 
their mesial ends, their shafts being straight and delicate; and there 
are no pterapophysial processes; they meet the tympanies in sub- 
circular heads, of no great size, just below the orbital processes. These 
latter elements possess very broad and twisted mastoid prolongations, 
with the usual double facet and intervening depression below; and the 
orbital apophyses are pointed at their extremities^ sometimes slightly 
clubbed, beiug turned gently upwards. The segments composing the 
infraorbital bar have long since become one single bone, a slender style 
fulfilling its ordinary functions. A sub-elliptical sesamoid is found at 
its proximal end, between it and the tympanic. The orbital cavities are 
capacious, and well divided from the rhinal chambers by the broad, 
quadrate lacrymals on either side ; their vaults are concavely arched, 
and their posterior walls quite extensive, looking almost directly for- 
wards. The foramen for the exit of the first pair from the brain-case 
has run into one irregular aperture; but rarely joins, in the adult, with 
the elliptical foramina for the optic nerves below them. The orbital 
septum is never complete, a vacuity of greater or less extent occurring 
near the center of the plate. Laterally we observe shallow temporal 
fossaB above elongated openings to the otocrane, that look downwards, 
forwards, and outwards, standing out quite prominently from the side of 
the skull. The squamosals throw forwards and downwards horizontally 
flattened apophyses, which by the aid of smaller ones from the ali- 
sphenoids, help it to guide the temporal muscles to their points of 
insertion. A moderately marked " cerebellar prominence" is found at 
its usual site, above the foramen magnum behind ; but we have never 
observed the foramina, caused by bone thinning, to occur on either side 
of it. In removing the cranial vault, we find the various fossaB unusually 
well defined, ana bounded by sharp borders; the carotids enter by 
separate openings at the base of the "sella turcica," which latter has a 
deep notch, mesiad, in its posterior wall. 

In the recent cranium, the internal and external tables are separated 
by an interspace of a millimeter or more, that is sparsely filled in by 
diploic tissue; but upon examining skulls that have been kept for a 
long time, and consequently become thoroughly dried, we cannot 
distinguish between the two tablets; the diploic tissue has entirely 
disappeared, and the whole roof is extremely attenuated and flexible. 
We are not prepared to explain how this remarkable change comes 
about. 

The lipoid arch bears out its usual ornithic and oscine characteristics, 
and does not require any special description here, as the author intends 



shufeldt.] OSTEOLOGY OF LANIUS. 721 

to furnish a more elaborate description of the skull when he comes to 
touch upon the Vireonidce ; a faithful outline of this arch is given, 
however, from a superior view in Fig. 101 of the plate. 

Before the young of this Shrikehas left the nest the numerous elements 
of the lower jaw have become fused together, so that during maceration 
the two rami rarely separate at any other point except the symphysis 
between the dentary elements. In the old bird it is a stout and strong 
bone, with sharp-pointed extremity beyond, and deeply scooped-out 
articular ends posteriorly, with blunted processes behind, and up-turned 
ones looking towards each other, mesiad. Externally the u sides of the 
jaw" are concave for their posterior two-thirds, and exhibit the usual 
elliptical foramen (Fig. 102) ; while the superior ramal borders are 
rounded and rise into slight prominences at the junction of the outer 
and middle thirds. As to the sense capsules, we find that the sclerotals 
are well developed and very accurately matched together; the usual 
ossicles of the organ of hearing likewise ossify. 

There are thirteen vertebrae devoted to the cervical portion of the 
spine, and, although they make a faiut attempt towards a raptorial ap- 
pearance, they are more oscine in their character than anything else, and 
are not noted for the prominence of any of their outstanding processes ; 
d sregarding the atlas, the first four bear neural spines, this feature not 
showing itself again until we find it in the last two, the thirteenth posess- 
ing it as well developed as any of the dorsals. The post- and prezyga- 
pophyses are markedly short, thus bringing all the segments quite near 
together, giving considerable stability to this division of the column. 
The parapophyses are very delicate where they are produced anteriorly 
at mid-neck, and quite inconspicuous above; the first four and the last 
six vertebrae bear hypapophyses, they being three-pronged on the last 
two; this limits the carotid canal to the fifth, sixth, and seventh cervi- 
cals, unusually slight protection for this important arterial branch. The 
vertebral canal commences in the tenth — i. e., in this segment it is com- 
pletely surrounded by bone, and continues its course through the axis; 
the last two or three vertebrae are very broad from side to side, the ulti- 
mate one bearing a free pair of vertebral ribs that have in their turn 
distinct uncinate processes. 

The neural tube as found in this section of the spine commences and 
terminates broadly and transversely elliptical, merging into the sub- 
circular as it nears its mid-portion at the middle of the neck; it is of 
considerable caliber throughout. The dorsal division of the spine has 
allotted to it five vertebrae, closely locked together, yet easily detached 
by ordinary maceration; their combined neural spines form one con- 
tinuous quadrate crest. These are fastened together above by the 
"arrow- head "joint that we have describedin other papers. There is very 
little difference in the lengths of the transverse processes, from first to 
last, so we do not find much change in the processes of the ribs they 
sustain, as to length of pedicles and tubercula. Short metapophysial 
ridges are found above the transverse processes; they never seem to 
attain sufficient length to connect the vertebral segments, however. 

The neural canal commences transversely elliptical, to terminate, much 
diminished in caliber, in the subcircular form. In the first dorsal we 
find a thin quadrate lamina of bone, projecting downwards and forwards 
in the mesial plane, as a well-developed hypapophysis ; the second sup- 
ports the merest apology for this process, and the remaining dorsals have 
none at all, though by compression of the centra a low ridge presents 
itself along their middles, which is only faintly perceptible in the last. 
There is a free pair of ribs for each dorsal vertebra, and these are mov- 



722 GEOLOGICAL SURVEY OF THE TERRITORIES. 

ably connected with the sternum by corresponding pairs of costal ribs, 
the whole structure and appearance being distinctly oscine in char- 
acter. The vertebral ribs are very slender below and not so much ex- 
panded above as they are in some other birds. Commencing with the 
last cervical rib, and continuing entirely through the dorsals, we find the 
series of epi pleural appendages complete on either side, and freely artic- 
ulated with the posterior borders of the ribs, with which they make an 
angle of about 45°, and attain sufficient length to overlap the rib in their 
immediate rear, though often in younger birds, and even some old ones, the 
last uncinate process does not reach the free sacral rib. The sternal ribs 
are quite delicately fashioned, and support, as usual, anteriorly the trans- 
verse heads for articulation with the costal border of the sternum, while 
posteriorly we discern the moderately upturned and clubbed extremities 
with shallow facettes for the inferior ends of the vertebral ribs. 

The sternum of the Loggerhead Shrike is almost or quite typically 
"cantorial" in its outlines, but only feebly pre-eminent in those features 
that stamp it as belonging to a bird of any great power of flight. The 
manubrium, directed upwards and forwards, springs from a solid base to 
become bifurcate at its anterior extremity and throw down a sharp 
border below, that becomes continuous with the carinal margin in 
front; the coracoidal grooves pass round laterally well beneath the 
costal processes, and merge into each other, mesiad, their point of meet- 
ing being denoted by an elliptical depression, at the base of which we 
occasionally find a single pneumatic foramen. The costal processes rear 
themselves upwards, forwards, and outwards, being broad but thin lam- 
inae of bone, impressed upon their posterior margins by the five trans- 
verse facettes for the sternal ribs. The " body " is concave above, sloping 
to a shallow, osseous gutter, lying in the mesial plane directly over the 
keel; beyond, in this groove we observe a few scattered foramina for 
the admission of air to the more solid structures of this confluent haemal 
spine. Behind, the bone is one-notched on either side, cutting out lat- 
eral processes with expanded posterior ends and a broad mid-xiphoid al 
portion — the direct continuation of the sternal body — (Fig. 92). The 
;; carina" below averages about 7 millimeters at its deepest part; ante- 
riorly it protrudes as a rounded carinal angle, from which point its inferior 
boundary sweeps backwards by a gentle convex curve to terminate in 
a diminutive triangular space at the middle of the xiphoidal process 
beneath. 

The sides of the keel present for examination well-defined subcostal, 
pectoral, and carinal ridges; the latter falls on either side from the base 
of the manubrial process to near the carinal angle, just within the bor- 
der, and sometimes has a thickened backward branch starting trom its 
lower end. The confluent pelvis, in common with the majority of pas- 
serine birds, has that strikingly angular outline, due largely to sharpened 
borders and outstanding spiny processes. There are ten vertebrae in 
its " sacrum," all unusually firmly fused together, vacuities only occa- 
sionally occurring among the diapophyses of the ultimate few, three or 
four at most. The pre-acetabular region of the ilia on either side is 
deeply concave, this concavity being carried up over the anti-trochauters 
to terminate in shallow grooves over tue ischiatic foramina. The greater 
share of this surface looks almost directly outwards and only slightly 
upwards. The ilio-neural canals are divided by the confluent spines of 
the first four or five vertebrae, they vary in width in different individuals, 
and terminate at points opposite the cotyloid cavities, at which point 
the neural spine suddenly becomes compressed, or rather annihilated, and 
the sacrum sustains a flattened surface to the ultimate boundary of the 



8HUFELDT.] OSTEOLOGY OF LANIUS. 723 

bone. The post-acetabtilar regions are of about one-third the extent of 
the surfaces anterior to the cotyloid rings ; they are produced behind in 
strong and clubbed processes, the outer margins of which are the termi- 
nations of the gluteal ridges or lines continuous with these ridges ; these 
surfaces are convex and narrowed by the encroachment of the broad 
sacrum (Fig. 103). 

Laterally the ilium overhangs the extensive and elliptical ischiatic 
foramen, which is bounded in front by the anti- trochanter, directed back- 
wards and slightly outwards ; the cotyloid ring is circular and but little 
difference exists between the diameters of its inner and outer peripheries. 
The obdurator foramen is very small and varies in the figure of its out- 
line, though generally assuming more or less the form of the ellipse ; 
the broad and thin hinder blade of the ischium again dips down to meet 
the slender pubic shaft, just before its' termination, to shut in an elon- 
gated spindle-shaped tendinal vacuity. (Fig. 100.) 

Upon the ventral aspect of the pelvis, we note that the bone affords 
no shelter whatever for the important organs it incloses until we pass 
the fourth sacral segment and the very decided vertebral swelling to 
form the sinus rhomboidalis ; it then drops into a deep depression on 
either side, whose concavities and convexities correspond with tHose de- 
scribed and attributed to the dorsal surface. The apophysial braces 
thrown out by the vertebrae are extremely slender, except in the cases 
of the first and fourth : the former segment bears a free pair of slender 
pleurapophyses, whose haemapophyses articulate along the posterior 
border of the ultimate sternal ribs, as do some of the inferior so-called 
"costal cartilages" in anthropotomy, lacking the necessary length to ar- 
rive at the costal borders of the great ventral haemal spine, constituting 
a common ornithic character. These sacral ribs rarely or never support 
uncinate processes. 

Six segments are devoted, in this Shrike, to the coccygeal division of 
the column, exclusive of the pygostyle ; they share the same fate, with 
their fellows and representatives in nearly all of the class Aves, in hav- 
ing many of their original vertebral components either rudimentary or 
entirely suppressed ; the neural spines, hooking oyer each other, ante- 
riorly, become more and more feebly developed as we proceed backwards ; 
this order of things is just reversed when we come to examine the hypa- 
pophyses on the nether aspects. The neural canal that passes through 
them dwindles to mere capillary dimensions before reaching the pygo- 
style, into which bone it enters but a very short distance. 

The transverse processes of the caudal vertebras are bent downwards, 
compressed horizontally, broad, and show but slight differences in length, 
before reaching the last one, in which they are shorter. The lamina of 
the pygostyle has the outline of an isosceles triangle, being truncate at 
its apex; the 'body" below is of a substantial structure, barely dilated 
behind, and otherwise presenting the usual characteristics as found 
among the oscines. 

The bones of the scapular arcli are all free and independent of each 
other, the stability of their relative position depending upon strong lig- 
aments in the living bird. The blade of the scapula is quite narrow, and, 
in the vast majority of cases, extends across the dorsal pleurapophyses, 
or vertebral ribs, its distal end being obliquely truncate, from within, out- 
wards; the blade-like portion is brought, up in close juxtaposition with 
that portion of the bone that affords the scapular moiety of the glenoid 
fossa. Its acromial x>rocess is very short, owing to the fact that it has 
to proceed but a short way before it abuts against the much -expanded 
head of clavicle, on either side; it forms with the coracoid the usual ten- 



724 GEOLOGICAL SURVEY OF THE TERRITORIES. 

dinal canal between the two bones. The head of the coracoid rears 
well above the glenoid cavity, in order to afford the required surface 
npon its mesial aspect for the broad clavicular extremity that rests 
against it; upon its opposite side it offers the usual surface to assist in 
completing the cavity for head of humerus. The shaft of the bone is 
very slender and cylindrical for its major part, and the wing-like exten- 
sion, so broad in many birds, is here but a meager osseous scale attached 
to the side of the shaft, for its outer and lower half, becoming continuous 
with the formal dilatation of the bone below; for the sternal articulation, 
this is transversely concave and very narrow. 

The minute pneumatic perforations of the scapulae and coracoids oc- 
cupy their usual sites back of the glenoid cavity, under the protection 
of the tendinal canal, at the heads of the bones. The united clavicles, 
or the furcuhun, inclines decidedly to the U-shaped variety (Figs. 94, 
95); we have already alluded to the fact as how broad, yet compressed, 
their scapular ends are found to be; from these heads the shaft-like 
portions fall downward, with a gentle curve backward to meet and sup- 
port the mesial and usual clavicular lamina, which here lies in that re- 
cess formed by the anterior and concave border of the carina of the 
sternum (Fig. 100). 

Directing our attention again to the shoulder-joint we discover that 
this Shrike is another example of those birds in which that little peg- 
like ossicle, the os humero-scapulare is found, here attached by its usual 
ligaments to the upper and back part of the articulation and fulfilling 
its ordinary function. The humerus of Lanius bears the closest resem- 
blance to that bone as found in many of the family Turdidce — particu- 
larly does this apply to Mimas polyglottus, a bird the Shrike not un- 
successfully apes in point of external coloration. 

The head, in most individuals, is well bent, anconal, and supports a 
short radial and not lofty crest, with the usual ulnar tuberosity over- 
hanging an ample pneumatic fossa. The shaft is quite straight and 
nearly cylindrical, its distal and expanded extremity presenting quite a 
unique appearance (Figs. 96 and 97). The internal and external con- 
dyles are distinct processes, the former projecting almost directly back- 
wards, the latter forwards and upwards; the olecranon fossa is likewise 
clearly defined, and on the palmar aspect we observe the oblique and 
ulnar facets unusually prominent. The humerus is the only bone of the 
pectoral limb that has air admitted to its interior, the bones of the anti- 
brachium and pinion lacking this rather rare prerogative. 

The ulna is more than four times the bulk of the radius, being, as in 
most vertebrates, the main support of the forearm; there is scarcely any 
perceptible curvature along its well-balanced and cylindrical shaft,Avhich 
presents a row of distinct little tubercles for the bases of the quills of the 
secondaries. Its proximal end presents for examination a prominent 
olecranon process, directed backwards, and the greater and lesser sig- 
moid cavities on its anconal aspect; the distal extremity is rather under 
the average in point of eminence, but shows all the usual indentations 
and surfaces to accommodate this end of the bone to the wrist and radius. 
The radius differs principally in having a general curvature distributed 
along its subtrihedral shaft, rather than having it confined to its proximal 
thiicl, as in many birds; otherwise it presents its ordinary ornithic char- 
acteristics. 

Among the mature birds representing the Laniidcc, as in so many 
other families, the carpus is composed of the two free ossicles, the cunei- 
form and the scapholunar, which are here impressed by their usual ar- 
ticulating f-icettes, for the radial and ulna trochlea? and the metacarpus, 



6HUFELDT.] OSTEOLOGY OF LANIUS. 725 

and although we have the young of this Shrike before us, the limits of 
this paper will not allow a critical description of this interesting and 
important region of the skeleton, that can only be obtained by careful 
study of the youngling. 

The manus contains its customary complement of bonelets, as seen in 
the pinions of the major part of the class in general (Fig. 100) ; medius 
and annularis metacarpals are firmly united together, and with the short 
first metacarpal that bears the index; the broad phalanx of the second 
is concave upon its anconal aspect and supports below the distal joint 
of the hand; the smallest phalanx of all is freely attached to annularis, 
which latter metacarpal extends some little distance below its stouter 
fellow, the medius. The pelvic limb is non-pneumatic, and consists in 
the adults of the usual number of bones, the patella being present. 
The femur, less than 2.5 centimeters long, has no trochanter minor, and 
the larger process of this name is but feebly produced ; the head, with 
its single depression for the ligamentum teres may justly be said to 
be at right angles with the cylindrical shaft, which latter is slightly 
convex forwards ; the condyles are well developed and the outer one pre- 
sents the usual fibular groove. The tibia presents nothing that differs 
in any marked extent from the oscines in general; it has no rotular 
process, but the pro-and ecto-cnemial apophyses are well produced and 
turned slightly outwards; at its distal end we observe, anteriorly, the 
usual tendinal bony bridge for extensor tendons. The fibula can be 
detached from the tibia by maceration, but its lower extremity spins out 
into a mere thread at the junction of mid and lower thirds of the latter bone. 
There are no free tarsal segments, and the same remarks apply here as 
we used in speaking of the wrist-joint above. The tarso-metatarsus 
(Fig. 99) is very delicately constructed below, while above it is stouter 
and presents immediately back of its head the process we have called ten- 
dinous, pierced by two pairsof foramina. A thin lamina of bone extends 
along its shaft behind. We observe that the os metatarsale accessorium is 
unusually large, as is the toe it supports; but otherwise the internodes 
are arranged upon the general plan of the oscine loot, which brings to 
our mind nothing of a raptorial type, except, perhaps, as we lenow the 
bird, the decided curvature of the hind claw, which is still further 
ncreased and lengthened when armed with its horny theea. 



726 GEOLOGICAL SURVEY OF THE TEKRITOEIES. 



PLATE XIV. 

Fig. 92. — The sTemum. from below. Lanius ludoriciamt* excHdf(oride9~ 
Fig. 93. — Right scapula and coracoid. outer aspect. 
Fig. 94. — Clavicular arch, from in front. 
Fig. 95. — Head of clavicle, right limb, outer aspect. 
Fig. 9*3. — Eight humerus, palmar aspect. 
Fig. 97. — The same, anconal aspect. 
Fig. RL — Eight femnr. posterior aspect. 
Fig. M9. — Left tarso-metatarsns. anterior aspeet. 

Fig. WO. — Skeleton of adult £, Lamms fttdo'.'icianm exeiiNloriifes ; the left free ver- 
tebral and costal ribs and the pectoral limb have been removed. 
Fig. 101. — The hyoid arch, from above. 
Fig, 102. — The lower mandible, from above. 
Fig. 103. — The pelvis, from above 
Fig. 10L — Superior aspect of skuLL the lo.\rer mandible having, been remaved. 



OSTEOLOGY OF THE CATHARTIDjE. 
By K. W. Shttfeldt, M. D. 

Captain, Medical Department, U. S. Army. 



Notwithstanding the fact that living birds, taken as a class, form one 
of the best isolated groups we have in nature, their arrangement into 
families and the division of these families into genera is still on a very 
unsatisfactory basis. This, no doubt, has been largely due to the fact 
that all of the variously proposed classifications have rested almost ex- 
clusively upon external characters; only an author here and there point- 
ing out in his writings differences in internal structure. Of recent years, 
however, this imperfect system and highly uncertain course has been 
very much modified and improved by the attention given to avian anat- 
omy by quite a number of earnest and painstaking zootomists. The 
published labors of these gentlemen and the facts they have succeeded 
in elucidating have already made their impression upon the problem to 
be solved, and. in man3 r cases the family lines are now more sharply 
defined. 

We must all agree, though, that this good work is yet in its infancy, 
and that the question of the classification of birds can only be definitely 
se.tled when we are thoroughly familiar with the habits, external 
structure, and characteristics of each species, and these are combined 
with a complete knowledge of their internal anatomy in all of its minutest 
details. The classification resting upon the digested and compared 
facts of such a knowledge as this can be the only perfect one, and the 
only one that will not be subject to sudden surprises and consequent 
changes, rendered necessary by periodical discoveries in structure and 
the development of new anatomical facts. In short, as distant as the 
day may seem, we will only be able to devise a perfect classification of 
living objects in nature when we have mastered their anatomy, using 
the word in its broadest sense. So that well-established and recorded 
anatomical data become particularly valuable to descriptive zoologists, 
and we feel sure that it is the aim of every one interested in this im- 
portant subject to further the efforts of those now engaged in this par- 
ticular branch of the science, in anyway in their power, in order to 
meet the end in view. Probably there is no better illustration afforded 
us anywhere in the recent literature of the science of ornithology, in 
which anatomy has played a more prominent part in deciding classifi- 
catory division, than in the separation of the Cathartidce from the Old 
World Vultures, nor in none, if it had continued to be relied upon, where 
simple external characteristics could have been more misleading. In 
this particular instance the osseous system was the one that finally ren- 
dered the principal assistance in determining the dividing line. 

Accepting as we must, then, these prerequisites to a sound classifica- 
tion, how idle it is for any one to attempt to define permanent family 
lines that will stand the test of time and research by any single set of 

727 



728 geological survey of the teeeitob:i> 

characters: as. for instance, cranial peculiarities, or such forms as the 
sternum may assume. It seems to me that this latter bone would be 
particularly unreliable to adopt for any such purpose, for we will soon 

: in the Cathartidce that its shape appears to vary with the age of 
the luaL, and a description of the bone in one bird, apparently 

an adult, would not answer for another of the same species, and per- 
haps of the same, or nearly th j same, age. 

At rhe present writing it is well _reed among ornithologists, and, 
•rrectly so. that The Gatkartidee, or American Vultures, 
form a family selves listinct :he vulturine Falconidw 

of the Old World. This rather recent step, taken as it was through the 
aid of modern dies in , will, in all probability, be a final 

one, ami the Vultures of our continent will, we ^iuk. always be con- 
as forming a well-defined family to which we can now hardly 
1c >fc for any new additions. The writer trusts that in the present 
paper he will be able \ lemoustrate to his readers, by the study of the 
skeletons : these birds, th eological characters of the group, 

me of which are new, . still furthei tow -taining tb~ - -._ 

made and refer:- te >v rhe [uestion as to how th jirtidm 

_ juld be divide _ ericallyis quite another thing, and its discussion 
will be postponed until we ha v ined the various skeletons of the 

membe: - of the family and compared our data with our present knowl- 
i of the other systems economy. 

Itures, as a rule, are found in the warmer climates, where they 
pursue their useful as s veugers, feeding upon all carrion 

that chance may th: :beir way. To carry on :ode of living 

find certain mark :- tui - presenting themselves to us in their 
superficial anatomy or external topography that are common to all of 
them. The m<>st important of these | nuts we ~ill hastily -view here, 
b :hat the reader may have them at hand as ex'trinal landmarks or 
refei ences t vhere comparison becomes possible, with points 

upon the skeleton within. In the fi: 8, we find that their upper 

mandibles terminate anteriorly in a well developed hook, which is much 
"-nned when encased in its horny covering: in fact, it requires 
the addition of this integumental sheath to approximate the mandibu- 
lar margins in any of the Co.:hartida: . The head and a greater or less 
ex: upper third of the ne : is devoid of feathers, a fine down 

ing found only in. the young .-.:■.'.-. I - : s no osseous septum na- 
rinm present, and the externa] nas 1 apertures or nostrils are 

made to assume different degrees of perviousm — the surrounding 
r. They have an external cla^ - horn, as in the ungual 

joints of the :—:. that articulates by its th the apex of the first 

digit in manus. A web is fouud between the inner and middle and mid- 
dle and outer toes occupying more or less of the basal thirds. Cla^> 
of the feet large, strong and curved, though these members are n 
fashioned for grasping purpos we elieve Vultures carry 

their plunder away by means of their feet . the — ■ .: ~ed 

claws rather to hold down firmly the portion of the carcass upon which 
they may be feeding while they tear it in shreds by their powerful 
hooked beaks. It will be recognized at once hat some o harac- 

- just enumerated have nothing whatever to do with the present 
mode of living of these . but may have done so in their am-it-nt 

anc-- - e refea particularly to the externa] claw exhibited on t 

manus. a character thev possess in common wirh some, if not all. of the 
Old World Vultn: 

The family Gatkartida^ as defined by its a st recent d— ::s. con- 



shufelot.] OSTEOLOGY OF THE CATHARTLTLE. 729 

sists of the following genera and species : First. Sarcorliamphiis, con- 
taining 8. gryphus, the Condor of the Andes, a bird whose range is so 
well known as not to require any definition from ine here. Gyparchus 
papa, the King Vulture, a species which, until recently, was supposed 
never to enter the confines of the United States, but inhabited the sub- 
tropical countries to our southward. Its presence at the present writ- 
ing, however, is strongly suspected in the Territory of Arizona, as refer- 
red to by Dr. Coues in the Bulletin of the Nuttall Ornithological Club 
for October, 1881, under General Notes. Third. Pseudogryphus, contain- 
ing the single species P. calif ornianus, the Californian Condor, a huge 
Vulture that is confined to our western coasts and principally to the 
State from w T hich it derives its name. Fourth. Cathartes, containing 
C. aura, C. burrovianus and G. pernigra. The range of the first of these, 
the common Turkey Buzzard, is thus defined by Mr. Eidgway : 

"Probably none of the birds of America have so extended a distri- 
bution as this Vulture, occurring as it does in greater or less abundance 
from high northern latitudes at the Saskatchewan, throughout North 
America, from the Atlantic to the Pacific, and in all portions of South 
America, even to the Straits of Magellan. (Hist, of N. A. Birds; Bd., 
Brewer & Ridw., pp. 345.)" 

From my own experience it can be said that this Vulture was found 
to be quite common during the years 1877-'80 throughout the northern 
territorial districts of the United States. 

In April, 1880, Ridgway calls our attention to 0. burrovianus and C. 
pernigra in the following words (Nutt. Ornith. Bull., Apr., 1880, p. 83): 

Cathartes burrovianns, Cass. — Recent authorities* having almost uniformly ignored 
the claims of this bird to specific rank, I have, in the absence of any opportunity to 
examine the type specimen in the museum of the Philadelphia Academy, carefully 
read Mr. Cassin's description in order to satisfy myself whether we are justified in the 
suspicion that Mr. Cassin's supposed species was based on a small specimen of C. aura. 
Upon reading Mr. Cassin's description I was surprised to find how ^ell and unmis- 
takably it applied to the bird usually called C. urubitinga Pelz. in every particular. 
In the description, as quoted below, I have italicized the phrases which are strictly 
and peculiarly diagnostic of C. urubitinga, in order to show at a glance how certain it 
is that Cassin's C, burrovianus is the same bird. The only question, it appears to me, 
can be as to the locality, which may be erroneous, since C. urubitinga is not known to 
occur anywhere out of Eastern South America, though the evidence to this effect, it 
should be remembered, is purely negative. 

The earliest notice of this species is that of Brisson (1760), the Vultur brasiliensis of 
this author being unquestionably the same species, as his full and very accurate descrip- 
tion clearly shows. Therefore, it is quite possible that some author may have applied 
the name brasiliensis to the species under consideration before Mr. Cassin's name &«?•- 
rovianus was bestowed upon it, in which event the proper specific term would be 
brasiliensis, and not burrovianus. I cannot rind, however, that such use of Brisson's 
name has been made. It is altogether probable that burrovianus will stand. 

Mr. Cassin's description (Pr. Philad. Acad., March, 1845, p. 212) is as follows: 

".Head naked, smooth, with the nostrils large and oval ; plumage of the body entirely 
black, with a greenish-blue gloss, paler beneath ; the feathers extend upwards on the 
back of the neck ; a small bare space on the breast. Wings long, the quills and tail- 
feathers black, with the shafts of the primaries white and conspicuous; third primary 
largest. The smallest American Vulture known. 

" Total length (of skin) 22 inches, bill 2$, wing 18, tail 8£. 

Hab. — Near Vera Cruz. 

"This species resembles C. aura, Linn., in the shape of the bill and the nostrils, and 
in having the tail rounded, but differs from it not only in size, but the feathers extend 
upwards on the back of the neck and lie flat instead of forming a ruff ; the plumage of the 
specimen now described is black, none of the feathers having pale margins, as is commonly 
the case in specimens of C. aura ; the shafts of the primaries are clear white, and the 
head is more entirely destitute of downy feathers. The tarsi are longer and more 
slender. 

* Conf. Elliot, Illustr. Am. B., II, 18G6 ; Allen, Bull. Mus. Comp. Zool., ii, 1871, p. 
311; Sharpe, Cat. Ace. Brit. Mus., i, 1874, p. 28; Gurney, The Ibis, 1875, p. 94. 



730 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



"The head of C. burrovianus is quite smooth, in which, as in other respects, it is 
very different from C. atratus, Wilson. 

"This new species was obtained in the vicinity of Vera Cruz, by the late M. Bur- 
rough, M. D., in honor of whom I have named it, as a slight acknowledgment for his 
very valuable services to natural history and to this academy." 

Cathartes pernigra, Sharpe. — A specimen of this species is in the Maximilian collec- 
tion, at the American Museum, New York. It appears quite distinct from both C. 
aura and C. burrovianus, being, in fact, somewhat intermediate between the two. In 
size it is nearly, if not quite, equal to C. aura, and, like the latter, has the nape en- 
tirely bare of feathers, the plumage commencing abruptly about half way down the 
neck. The shafts of the primaries are a lighter brown than in C. aura, but not so 
white as in burrovianus. In regard to the plumage, however, there is a much closer re- 
semblance to C. burrovianus, the back and wings being wholly black, like the lower 
parts, without a trace of the light-brownish borders to the feathers, so conspicuous in 
aura. The black is also much less glossy than in the latter. 

The specimen (male) is, unfortunately, not quite adult, the bill being partly black- 
ish, and the nape covered with a soft dusky down. The measurements are as follows: 
Wing, 20.00; tail, 12.00; culrnen (chord of the arch), .85; tarsus, 2.50; middle toe, 
2.40. 

The bill and feet appear more slender than those of C. aura. 

In conversation with Mr. Kidgwny at the present date (Xovember 
12, 1881) we find him still adhering to the views he so clearly set forth 
in the Bulletin the year before, still declaring that C. burrovianus and 
C. pernigra are both " good species." 

We had the opportunity to examine a mounted specimen of the former 
in the cabinet of Vultures at the Smithsonian Institution on the same 
day, and must agree with Mr. Ridgway, that as far as external charac- 
ters are concerned, as compared with aura, the birds certainly appear 
distinct. We observed the additional fact that the nostrils in burrovi- 
anus were far more pervious than those in aura even 

A specimen of C. pernigra we never have had the good fortune to 
examine ; in fact, both of these Vultures are uncommonly rare in col- 
lections, and *he writer has thus intentionally takeu the pains to bring 
out all of the data that he has at his command bearing upon these two 
birds, as he must here confess to his reader that it has been found im- 
possible to secure a skeleton of 
either burrovianus or pernigra, 
so that the characteristics of the 
genus Cathartes must be drawn 
from its common representative, 
aura. We do not know of a 
skeleton of either burrovianus or 
pernigra being in any of the oste- 
ological cabinets in the United 
States, and it must be left to some 
more fortunate ornithotomist to 
compare the osteological charac- 
ters of these two birds, with what 
the author hopes to bring out in 
regard to the genus in which 
have been placed, from their 
northern representative. 




Pseudogryphus californanus. 



Fourth, and lastly, we have the genus Catharista, containing the single 
species C. atrata, the Carrion Crow, or Black Vulture. This bird is con- 
fined more or less to our Southern States, inhabiting particularly the 
maritime districts, and to various localities in South America, its habits 
and tastes always being those of a true Vulture, which it is. It will be 
remembered that for a long time many authors placed all of our ^sorth 
American Vultures in the one genus Cathartes; the group containing, 



shufeldt.] OSTEOLOGY OF THE CATHARTID^E. 731 

according to Coues' Key of North American Birds (1872), Cathartes cali- 
fornianus, C. aura, and C. atratus. They appear in Mr. Bidgway's 
Nomenclature of North American Birds (Bull. U. S. Nat. Museum, No. 
21, 1881) as we have given them above (G. papa not being included), 
viz, Pseudogryphus califomianus, Cathartes aura, and Catharista atrata.* 
We shall soon see, when the skeletology of these have been compared, 
how far the osteological characters will support this generic division. 

Before passing, however, to the examination of the skeletons of these 
birds, the author proposes to present his reader with a short table 
showing additional points in external structure that have been gleaned 
from various sources and authorities as well as from his own memo- 
randa. We feel sure that although in a paper proposing, as its title 
indicates, to be purely osteological in character, such a diversion will 
hardly be considered amiss. 

* Dr. Coues, in his Check List and Ornithological Dictionary of 1882, also adopts 
this arrangement. 



732 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



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SHUFELDT.] 



OSTEOLOGY OF THE CATHARTID^. 



733 




Cathartes burrcvianus. 




Cathartes aura. 



Many of the characters noted in the above table are from those ob- 
served by John Henry Gnrney (Descrip. Cat. of Eap. Birds in Norfolk 
and Norwich Mus., Loud., 1864), others are from Mr. Ridgway's Falconidce 
and Catharticlce (Hist, of N. A. B. by B., Br., 
and Bidgw.). This latter ornithologist de- 
scribes still one other Vulture found in South 
America (Nutt. Ornith. Bull., Apr., 1880), but 
it seems to us that it is not yet sufficiently 
proven that this bird is not 8. gryphus in im- 
mature plumage. Particularly may this be 
the case, as we do not positively know how 
long the Condor requires to attain the adult 
plumage, and, furthermore, we can hardly be- 
lieve that a bird of the size of the oue described 
could have so long escaped observation. However, in case we may 
be wrong in the matter, and future developments prove this to be a 
new Condor, we will here insert Mr. Bidgway's notes 
upon this Vulture, taken from the bulletin above 
quoted. 

Sarcorhamphus cequatorialis, Sharpe? — In the vivarium at Cen- 
tral Park, New York City, I saw in December, 1878, a Condor of 
uniform brown plumage, which Mr. Conklin, the director of the 
menagerie, informed me had been received July 23, 1875, and that 
it was three months old when captured. It was obtained on 
Mouut Conquences, Chili, and was presented by Eear-Admiral 
Collins, U. S. N. The fact that this example had not yet, when 
nearly four years old, begun to assume the plumage of S. gi'yphus, jtroves conclusively 
either that the latter species retains the livery of the young until four or more years 
of age, or that there really is, as has been asserted by authors, a sjiecies of Condor 
among the Andes which has permanently a uniform brown plumage, something like 
that of Gyps fulvus.* The locality of this specimen would extend considerably the 
range of S. cequatorialis, Mr. Sharpe giving only Ecuador, and doubtfully Colombia, as 
the habitat of his species. 

I will also remark here, and I have noted it in birds of various species, 
more especially in a living specimen of Haliaetus 
leucocephalus that I had in my possession for a long 
time, that birds do not seem to acquire their adult 
plumage nearly so soon when they are kept in a 
state of confinement as when leading their natural 
life. Other authors have likewise noted this fact. 
With so many curious cases on record among other 
species, it is hard telling exactly what effect such 
an entire change would have upon S.gryphus, hav- 
ing been taken young, removed to an entirely dif- 
ferent climate, kept confined, and no doubt hav- 
ing a radical chauge made in its diet. 

Before closing what we have to say upon the external characters of 

* Since the above was written, Mr. Conklin has favored me with the following par- 
ticulars, under date of February 10, 1880, in response to my inquiry as to the present 
condition of this specimen : "The plumage remains still unchanged, except that the 
ruff about the neck is somewhat fuller, and has a little sprinkle of white ihrough the 
down . . . It has not increased much since then (July 23, 1875, the time when received 
at the menagerie), either in size or weight. The bill is black at the base, the apical 
half ivory-white; head bare, no wattles; iris dark brown." Mr. Lawrence has also 
favored me with the following transcript from his note-book, April 1, 1876 : "Condor, 
said to be nine months old. Bill black ; cere and naked sides of head grayish-black; 
head sparsely covered with short downy feathers of a smoky black ; plumage, in gen- 
eral, of a dark snuff-brown." August, 1877: "No change except the development of 
the ruff, which is colored like the back." "The ruff is now (February 23, 1*80) more 
full, but no appeara; ce of becoming white ; underneath the feathers are whitish." 




Catharista atrata. 



73-4 GEOLOGICAL SURVEY OF THE TERRITORIES. 

these birds, the writer inserts the following valuable quotations from 
Nitzsch, an author who paid no little attention to the external charac- 
teristics of birds, more particularly the feather tracts. 

The chief pterylographic character of These, as of the Old World Vultures, is to he 
found in the formation of the pectoral portion of the iuferior tract. This is not sepa- 
rated by a space from the jugular portion, but the two sections of the tract are per- 
fectly continuous. Just as the inferior tract, gradually enlarging. Las arrived over 
the pectoral muscles, it receives the axillary tract, and thus acquires a very remark- 
able breadth. It then divides into two branrhes. which, however, are generally weak, 
and are rendered indistinct by the circumstance that the feathers of the tract are 
more scattered. The two branches are at hrst of equal width, and run parallel : to- 
wards the extremity of the great pectoral muscles they curve in towards each other; 
and whilst the inner one is continued unchanged as the ventral portion, the outer one 
is narrowed, and returns by a narrow process, running along the margin of the mus- 
culus pecto ralis major to the ventral part again. 

In this way the two branches inclose a pretty large insular space, the whole tract 
thus presenting very close resemblance to that of Cenlropus (refers to }>late). 
This, however, does not conclude the list of the pterylographic peculiarities of the 
American Vultures : but we must add : 1. The amalgamation of the jugular part of 
the inferior tract with the cervical portion of the spinal tract, which would convert 
the plumage of the lower part of the neck into a continuous one. if the space did not 
entirely or partially penetrate it. 2. The remarkable narrowing of the dorsal portion of 
the spinal tract, which frequently consists only of two rows of feathers, gradually 
diverging anteriorly, and connected with the branches of the fork of the cervical part. 
3. The presence of a large lumbar tract. 4. The constant and invariable presence of 
twelve feathers of the tail. 5. The absence of a circlet of feathers at the apex of th* 
oil-gland. The broad, obtuse form of this organ seems to stand in relation to this. At 
its extremity there are two distinct orinces. 6. The covering of the feet may also be 
cited as a characteristic element : it consists of small scales upon the tarsus, but of 
scutes upon the whole of the toes : moreover, besides the outer and middle toes, the 
middle and inner toes are united by a membrane. 

The elongated nostrils, parallel to the longitudinal axis of the beak, certainly remind 
one of Neophron ; but the absence of a bony septum between them is one of the most 
characteristic external distinctions of the American Vultures: the tongue, moreover, 
has a series of teeth on its margin, at least in C. papa and C. aura. 

I have examined C. gryphus. papa, aura, and uruber (fcetens Illig. ). and found in all 
some little differences in the form of the tract, but no characters from which I could 
justify their division into the genera Sareorrhamphus and Catharie*. — (Pteryl. Trans., 
from German Ed., by P. L. Sclater. Lond.. 1~67. p. 50.) 

From this point the author goes on to give, under "Cathartes," de- 
scriptions of the feather tracts of the birds that he mentions as having 
examined in the above list, in detail. 

The writer has to present to his reader the following material for ex- 
amination, of the skeletons of the Catliartida. upon which to base the 
results of his monograph. At the outstart we must deplore the fact 
that we do not possess the skeletons of the young of any of the family 
under consideration. To be sure, specimens of the young of Psevdo- 
gryphwi have been obtained on several occasions, but at the present 
writing there are none in the Smithsonian Institution, and should such 
be obtained at a later date they can be discussed in some future paper. 
Of Sarcorhamphm gryphus we have in our study one very good skeleton, 
the property of the Smithsonian Institution, and kindly lent us for the 
purpose in hand: it however lacks the osseous part of the claw of the 
hrst digit of the hand. This is also the case in our skeleton of Gypar- 
chus papa, an unusually fine skeleton belonging to the Army Medical 
Museum, prepared by Prof. H. A. Ward, of Rochester, and obtained in 
Ecuador, probably by some of his collectors. Of Pseudogryphus caJifor- 
nkvaus. also from the osteological cabinets of the Smithsonian Institution, 
we have two imperfect skeletons; they, however, show the cranium and 
the majority of the principal bones. We may consider ourselves fortu- 
nate, however, in having even this much, for this Vulture is becoming 
rarer and rarer each year that slips by, a fact that is to be accounted for 



shufeldt.J OSTEOLOGY OF THE CATHARTID.E. 735 

through many causes, chief among which is its destruction by poisoned 
flesh and carcasses thrown out by the cattle-owners of the countries it in- 
habits, to destroy the wolves and bears. Then, too, it no doubt, being a 
bird not difficult to approach, forms a good target for many a hunter who 
chances to meet it with rifle in hand, and happens to be ambitious to add 
to his record the fact of having slain the largest bird in North America. 
For Cathartes aura we have numerous skeletons and parts of skeletons 
obtained from various sources, some from the Smithsonian collection, 
others from the Army Medical Museum, and one or two from my own 
cabinet, obtained by myself while in the territorial districts of the inte- 
rior. We have already stated that we lack any osseous material from 
the two remaining species of this genus, C. burrovianus and C. pernigra, 
and are of the opinion that the skeletons of these birds are not to be 
found in any of the collections in this country. 

Lastly, Catharista atrata is represented by several good skeletons and 
many crania and separate bones, placed at our disposal by the institu- 
tions above mentioned, being amply sufficient for the purposes of de- 
scription and comparison with the other members of the family. In 
addition, we have before us a good skeleton of Gypogeranus serpenta- 
riuSj one very complete mouuted skeleton of Neophron percnopterus, 
from Abyssinia, from the collection of the Army Medical Museum, and 
finally a mass of odds and ends of skeletons of Hawks, Eagles, and Fal- 
cons, that bear more or less upon the discussion of such a subject as 
we have undertaken, the majority of which latter material we are deeply 
indebted to the kindness of Professor Baird, and the exertions of Mr. 
Eidgway, in the way of selection. 

In the matter of plates, we trust we have laid a sufficient number 
before our reader to thoroughly illustrate all we have to say in the text, 
and the author assures you that be has not spared his pains in choos- 
ing subjects such as he deemed would best fulfill the end in view. Per- 
haps Pseudogryplius in this choice has taken the lion's share, but we 
must remember that in extenuation of this claim our Californian Vul- 
ture will soon be reckoned among the birds that were, and life-size 
portraits of the bones of his skeleton will be ranked among those of the 
famed Dodo, and the great Auk. His cranium and the outline sketch 
of the metacarpus are from my own pencil, but the remainder of the 
bones of his skeleton are from photographs taken at the Arm}* Medical 
Museum, under my own supervision, and are all life-size. The skull of 8. 
gryphus was kindly drawn for me by Dr. J. 0. MeOonnell, of the Army 
Medical Museum, to whom science is so much indebted for illustrations 
in so many of its branches. In another plate we present an entire skel- 
eton of G. papa from a photograph and reduced to one-third of its size j 
this figure shows well the tout ensemble of the bones of the skeleton in one 
of our typical Vultures, as well as the relations of size and position of 
these bones as compared with each other. It is the Equaclor specimen 
referred to above. The two plates, figuring three views of the cranium 
of G. aura, with two views of its sternum and its furculum and remainder 
of scapular arch, are India ink productions of the writer's, being drawn 
from a fine specimen of this bird, secured by him in the Laramie range 
of hills in Wyoming. The remainder of the figures are life-size views 
of the subjects they represent, from photographs taken at the Army 
Medical Museum, and are very accurate conceptions of the bones they 
depict. Their detailed descriptions must, however, be deferred until 
we come to describe the parts of the various skeletons from which they 
were taken. 

Upon viewing the skeleton of one of these birds for the first time, 



736 GEOLOGICAL SURVEY OF THE TERRITORIES. 

such as we offer in Plate XV, fig. 105, G. papa, we are struck with the vul- 
turine appearance of the skull, with its more or less open, capacious ex- 
ternal nasal orifices, its hooked beak, its rather large orbital cavities 
and the good size of the bones at the bases of these orbits. The seg- 
ments of the vertebral column, or the vertebrae, are all free, from oc- 
ciput to sacrum, with their processes and arches well developed. We 
observe, too, the great length and size of the bones of the extremities ; 
particularly those of the brachium and anti-brachium of the upper, and 
the robust femora of the latter. To support these great limbs, we find 
these birds possess correspondingly well developed arches of the axial 
skeleton. ]Sor are the bones of the feet backward in indicating to us 
the avocation of their owners, attached as they are, too, to powerful 
leg bones. In short, the Cathartidce must be regarded as possessing 
skeletons composed of bones above the average, for the size of the 
bird, in point of volume and length. These characteristics, however, 
do not bring with them a correspondingly heavy skeleton, but quite 
the contrary, for if we pick up the bony framework of any of our Vul- 
tures we cannot help but being struck by its unusual lightness ; this is 
due to almost a universal pneumaticity of the bones composing it, and 
so marked a feature is this of the birds we are describing that it will 
not be considered out of place to give a general description of this 
condition here, so as to avoid, as much as possible, what would have 
to be a constant reference to the many pneumatic foramina that occur 
in nearly all of the bones that we shall come to describe further on. 

Let us choose a specimen of Catharista at rata that we have before us 
for this purpose, simply reminding the reader that at least some of 
these openings will necessarily have to be referred to again, wbile with 
others their description will cease here. In the skull of this Vulture, 
as in all others of the family, the only elements that seem to possess 
this property are the quadrate bones (the tympanies of my former 
papers), the posterior thirds of the limbs of the lower mandible, and 
the lacrymals. In the quadrate bones we usually find a single, rather 
large, sub-elliptical opening at the bases of the mastoid ends or pro- 
cesses with a large one at their summit inPseudogryphus — while several 
such foramina occur on the superior aspects of the articular extremities 
of the lower jaw 5 two of these latter seem to be constant and are found 
towards the points of their in-pointing prolongations. On the inner 
aspect of the lacrymals we find either one such pneumatic orifice or a 
group of several of them, and the air seems to attain complete access 
to these bones, and perhaps enters to a limited extent the wings of the 
ethmoid that they anchylose with on either side. It is very apparent 
that the walls of the brain-case or the bones that go to form the beak 
could not afford to make any such sacrifice of stability for the little 
additional lightness that would be gained thereby — the functions of 
these parts would evidently be weakened by it, affording as it would 
insufficient protection for the brain, and rendering the beak constantly 
liable to fracture. 

The author is free to confess that at the present writing he knows 
not of a single instance among the class where to look for a bird pos- 
sessing an atlas endowed with this property, and we find here in the 
Cathartidce again, this bone apparently solid, or at least absolutely non- 
pneumatic, while the following vertebra, the axis, is quite so. Consti- 
tuting, as its occurrence does in this bone, rather an uncommon condition 
for it, knowing as we do that in many birds where every vertebra will 
be found to be pneumatic, the atlantal and axial segments remain ex- 
empt. 



shufeldt.J OSTEOLOGY OF THE CATHAETID2E. 737 

In the cervical portion of the column one or more of these little aper- 
tures are found to pierce the centrum of each vertebra, within the ver- 
tebral canal or just outside of it, anteriorly. This is not a circumstance, 
however, we w T ill find when we come to examine the dorsal segments, for 
here about the bases of the diapophyses or transverse processes great 
circular openings occur that allow one a view directly through the ver- 
tebra, and the center of these segments are in some cases riddled with 
similar though smaller foramina. Such openings, likewise, are found to 
exist at the inferior aspects at the extremities of the diapophyses, and 
sometimes a few minute ones in the lnetapophysialoif-shoots beneath. 

The position of these foramina are but little changed, much less their 
number or size materially decreased or diminished in those vertebrae 
constituting the sacrum. After we pass the first three or lour segments 
of this division in C. aura, however, we seem to lose sight of them in 
the diapophyses, which is not the case among the species of the other 
genera, though they are very sparsely distributed in C. alrata. Arriv- 
ing at the coccygeal vertebras, we find that pneumatic foramina have 
disappeared entirely, and this holds good for all the Vultures and ap- 
plies equally well to the ultimate segment of this division, the pygo- 
style. We present two or three figures in which the extensive preii- 
matic openings that occur on the outer aspects of the expanded heads 
of the clavicles may be seen. In C. atrata, as in the others, these con- 
sist essentially, one each side, of a large cloaca that extends immedi- 
ately into the bone, rendering it one of the lightest for its size of any 
of the skeleton. About the entrance of this general opening we observe 
many smaller and circular or variously shaped ones irregularly arranged 
in groups, with sometimes only a single one beyond, isolated from the 
rest (Pseiidogryphus), or another separate little collection in this situa- 
tion ( C. aura). In the scapula the openings occur near the glenoid cavity, 
and again a grouplet beneath the ridge found at the crest of the acromial 
process, on its outer aspect, but the number or size of these foramina in 
this bone is never very great. 

In any member of the family, a careful search about the head of either 
coracoid is usually rewarded by the discovery of a few scattered open- 
ings of this sort, but these bones seem to depend almost entirely for the 
admission of air into their shafts and extremities upon a cluster of 
these foramina that are met in the recess of the dilated ends below, on 
their posterior aspects. 

Passing next to the sternum, we find in all the Vultures, less marked 
in C. atrata than any, a rounded and prominent ridge, extending back- 
wards from the anterior margin immediately posterior to the manubrial 
eminence, to the middle line of the concave and superior aspect of the 
body below. It is on either side of this ridge that we find pneumatic 
perforations that lead into the heavier portions of the bone, more particu- 
larly the promintory of the manubrium, and downwards into the thick- 
ened carinal ridge. Again, in the recesses above the subcostal ridges 
we discover irregular groups of these pneumatic foramina, that pass into 
the lateral portions of this bone; while lastly, the apertures through 
which additional aerification of the sternum takes place occur in the 
pits along the upper side of the costal borders, that we find among the 
facets of articulation for the sternal ribs. In these localities they have 
a tendency to gather around the bases of the eminences that support 
these facets, rather than be generally distributed throughout the depres- 
sions in question. (Plate XVII, fig. 107, Pseudogryphus.) 

In the free ribs, both those of the cervical division of the column, as 
well as the dorsal ones, we find these pneumatic foramina entering the 



738 GEOLOGICAL SURVEY OF THE TERRITORIES. 

bone as one large opening (C. aura) or by a group of several smaller 
ones (C. atrata) at the outer extremities of their necks, just below the 
tubercula, anteriorly. We have just said that they occur in the free ribs 
of the cervical portion of the spine; this is indeed the case, but of very 
rare occurrence, and we may almost add in perfect safety the ribs of the 
sacral vertebrae to the list of non-pneumatic bones of the Oathartidcp. 

The sternal ribs, at their inferior extremities, show like per I orations, 
more commonly on their anterior aspects, though by no means do we 
always find them wanting on the posterior faces; more likely would 
we be disappointed in our search for them in the last sternal ribs; and 
we doubt of their ever occurring in the set that articulate with the first 
pair of sacral ribs. 

The osseous portion of the basin, completed by the parial pelvic bones, 
is not usually permeated by air as thoroughly as it might be: indeed, in 
all those parts of the pelvis where the component bones become plate- 
like in character or flatten out, this property is denied them, and it is 
only where ridges become thickened from any cause whatever or other 
prominences take place that we fiud pneumatic foramina present. 
Notably these localities are three in number: Externally, in the recess 
on either side, just below the angle formed by the deflection of the glu- 
teal ridge, and above the antetrochanter. Internally, in two places 
about the cotyloid ring at the extremities of its horizontal diameter 
slightly produced, at a greater or less distance from its periphery, and 
again at the base of the posterior concavity formed by the ilium and 
ischium, immediately within the rounded margin of the ischiatic fora- 
men. 

Of the Vultures, G. atrata seems to possess the most pneumatic sacrum, 
while that of C. aura is the least so. In Pseudogryphus the foramina 
are particularly well supplied, forming good-sized groups at all the 
localities just specified. They are absent in the last situation mentioned, 
in C. aura, SarcorhampUus, and C. atrata, but in the last species quite 
a cluster occurs in the ilium on its internal surface, well within the upper 
margin of the ischiatic foramen. 

The pectoral limb of the Catliartidce is highly pneumatic from humerus 
to the last joint in manus, with the sole exception of the osseous core 
of the claw on the first digit. 

In the humerus the large pneumatic fossa, with the collection of 
smaller openings at its base on the anconal aspect of the head, is never 
absent ; in all but aura a few additional ones are found at this extrem- 
ity of the bone, on the opposite side, just beyond the articular surface. 
At the distal end of humerus other openiugs occur of no mean size. 
especially in G. papa and C. atrata. These are generally situated on 
the anconal aspect of the bone, in a shallow but circumscribed depres- 
sion that is found there, beyond the tubercles for articulation ; some- 
times a large one is found quite near the ulnar tubercle. 

The cubit and radius both have these little perforations well supplied 
them, especially about their proximal extremities, anconad; the number 
not being nearly so great at the other end of the bones. The two free 
carpal bones possess at least two or three of these foramina apiece, and 
we note on the anterior face of scapho-lunar in C. atrata two very siz- 
able ones, that seem quite constant for this species. 

In metacarpus, the bone is pierced by a single foramen just beyond 
os magnum, at the point where index and medius metacarpals joined in 
the growing bird. Other perf orations are found about both extremities 
of this segment in the angles and recesses formed by the conforma- 
tion given it by the amalgamation of second and third metacarpals. 



shufeldt.] OSTEOLOGY OF THE CATIIARTID^. 739 

The palmar aspect of all the long digits show one or more such aerial 
penetrations, while the blade-like expansion of the first phalanx of 
second metacarpal seems to be absolutely riddled by these foramina, 
lending to it a decidedly honeycomb appearance. 

The femur is the only bone of the pelvic limb that enjoys the condi- 
tion we are discussing. In it air gains access to its interior through a 
single opening {G. aura), or, as in most cases, several of them, on the 
anterior aspect of the bone, below the great trochanter, close to Che 
prolongation of its curling crest. In Fseudogryphus we discover a few 
additional ones on the upper face of the trochanter major, beyond the 
usual group. 

We may say, then, that as a rule for the Gathartidw, the only bones 
in their skeletons that do not receive air into their interiors through 
pueumatic foramina are certain portions of the cranium, lower maxilla 
and pelvis, the hyoid arch, the atlas, the coccygeal vertebras, including 
the pygostyle, the bones of the pelvic limb below the femur, and all sesa- 
moids and ossifications pertaining to the sense organs. 

Passing to the Falconidw and the Old World Vultures, we observe in 
the latter that a J though thej 7 have the greater share of their skeletons 
pueumatic, it is not nearly so perfect a condition as it has just been 
shown to be in our Gatharudw. In Neophron percnopterus the axis is 
non pneumatic, as are all the segments of the pectoral limb save the 
humerus. To these must be added the bones that we have just passed 
over and enumerated in the family in hand, as being so. 

We find the furculum in the Vulturinw a bone not nearly so well 
aerated as we have described it for the New World Vultures, it is still 
less so in Gypogeranus, and the same remarks will apply here in gen- 
eral for the remainder of the skeleton of this bird that we have this 
moment made in regard to the Vulturinw. In general they are equally 
applicable to the remainder of the family Falconidw. We believe that 
we will not go far astray in laying it down as a rule for the entire order 
Raptores, that this attribute of the skeleton seems to have arrived at 
its acme in Gatharista atrata, exists to a greater or less degree in all of 
the American Vultures, then becomes less marked as we pass to the 
vulturine Raptores of the Old World, among which we believe that the 
clavicles will be found to be far less pneumatic throughout, though the 
femora still remain so, down among the diurnal and nocturnal Raptores 
of both continents, until we arrive at such forms as Speotyto, where we 
know the entire pelvic limb to be exempt from this condition, and that 
air is excluded from more than half the bones of the skeleton of this 
ground-abiding Owl. 

Having dwelt now, and we trust with sufficient the roughness, upon 
the present classification of the Gathartidw, their leading and more 
important external similarities and differences, and then touched upon 
some of the more general characteristics of their skeletons, we will pro- 
ceed to investigate these latter more in detail, beginning, as we have 
done in former memoirs, with the skulls of the species to be treated. It 
is the object of the author during this investigation to keep these two 
things prominently before his reader, the first being the prime object of 
the work we have to do, viz, an accurate description of the skeleton of 
each species of the family as far as our material will allow us to give it ; 
and, secondly, from time to time, to compare the principal bones of 
these birds with those of members of their own family, and then with 
those of other families of near kin 5 our second aim being, so far as oste- 
ology will justify it, to test the all-iniportaut question as to whether 
the Gathartidw have been wisely arranged and distributed in the genera 



740 GEOLOGICAL SUL.VEY OF THE TERRITORIES. 

created for them, and. if such prove to be the case, to present the re- 
sults of our labor, and so apply it as to settle these birds in the places 
they now occupy upon a still firmer basis, to the end that we may have 
it said that classification has been assisted and furthered. 

Of the skull — It will be remembered that when we treated of the 
osteology of the North American Tetraomdee .Bull. U. S. Geol. and 
Geogr. Surv. Terr. Vol. YI, Xo. 2. pp. oil. and Plate Y, fig. 51) we 
made a careful dissection of the skull in the young bird before the seg- 
mental anchylosis commenced, and figured and named all the various 
bones that enter into the cranium of the majority of the class. At this 
date, however, the author does not advise the student to implicitly follow 
the nomenclature there given, nor accept all of the deductions set forth. 
We have already informed our reader of the fact that it will be impos- 
sible to carry this interesting part of the subject out in the case of the 
CathartidcB as we do not possess the necessary material. Still, the fig- 
ure above referred to. and the description given for the young of Centro- 
eercus. will no doubt assist us in approximating very closely the bound- 
aries and limits of many of the cranial bones, not that there is the 
slightest relationship among the species of the two families, i. e.. the 
let ra on idee and the CatJiartida\ but that certain rules hold good in each. 
The outward appearances, though, in one case is quite striking, and led 
Dr. Cones to say in his Key : 

••In a certain sense, they represent the gallinaceous type of structure : 
our species of Cathartes. for instance, bear a curious superficial resem- 
blance to a turkey." — Key to X. A. B., pp. 221. ed. 1872.) 

It is hardly necessary to say here that osteologically the species are 
quite distinct, as Meleagris has its skeleton unmistakably stamped with 
all the characteristics of the GaUince, as Catliartes aura has all those of 
the Yultures. the two being very different. 

In the crania of the Gathartidce we find that, after the different species 
have arrived at maturity, a union has taken place among the various seg- 
ments of the so-called vertebral arches that is quite above the average 
result or condition in birds generally, for upon examination we find that 
we have only as free bones the ossa quadrata, the pterygoids, bones of 
the sense capsules (except the ethmo-turbinals), the hyoidal segments of 
the tongue, and the inferior maxilla, all the others having firmly united 
together, very few sutural traces of original separation being left. 

Plates XXII and XXIY, figures 119 and 127. respectively, give good 
representations of the superior aspects of the skulls of G. aura and C. 
at rata. Many of the peculiarities we discern in these figures are com- 
mon also to the other members of the family. TVe are struck at once 
with the great breadth of the pseudo-articulation that goes generally 
by the name of the •• fronto-maxilliary." This is due principally to the 
fact that the nasal bones are unusually broad, in order to encircle the 
great sub-elliptical nostrils of these birds. This articulation, notwith- 
standing its width, is very mobile, and offers several departures from 
the usual mechanism of the system to which it belongs to interest us. 
We find that from the anterior border of the united laerymal, on either 
side, a niche exists that gives rise to a process, below which process. 
in turn, has its superior border grooved lengthwise and converted into 
au articular surface for a quadrate apophysis thrown out by the outer 
and posterior angle of either nasal, In a position of rest a foramen re- 
mains at the base of this niche in the laerymal formed by the two bones 
in question, allowing in action or movement the process of the nasal to 
glide into it. (PL XXII. fig. 118.) This is the state of things in G. aura. 

In Pseudogryphus the nasal process is much longer, cuivcd upwards, 



shufeldt.] OSTEOLOGY OF THE CATHALITID^E. 741 

and glides over a greater surface on the lacryraal. Tlio quadrate form 
of the nasal process becomes peg-like in atrata, really articulating in a 
socket on the interior aspect of the lacrymal. In the remaining species 
we find this joint more or less persistent, perhaps less so in the South 
American Condor. This mobility produces rather a confusing condi- 
tion at the pterygo-basi- sphenoidal articulation, for we have observed 
in all of the dry skulls of the Cathartidw that the pterapophysial pro- 
cesses of the basi-sphenoid never meet the facets on the pterygoids that 
are evidently intended for their articulation. This seems to be due to 
a warping upwards of the superior mandible during the process of dry- 
ing, drawing both the palatines and with them the pterygoids away 
from these pterapophyses. If we take the pains, however, to dissect 
the head of a recently-killed Vulture, as the author has done, we will 
at once appreciate the normal state of affairs, and find that by the slight- 
est pressure downwards of the upper bill the facets upon the pterygoids 
glide over the pterapophyses. We will find many of the illustrations 
representing them with the interspace between. Our own plates do so, 
and Professor Huxley has done so before us, as we find about 2 milli- 
meters of space between the pterygoids and the processes from the 
sphenoid in his view of the base of the cranium in C. aura, although he 
remarks in the text, "The basipterygoid processes are large and artic- 
ulate with the pterygoids." (Class, of B., Proc. Zool. Soc. Lond., 1867, 
pp. 440, fig. 22.) The reader will remember that in our figure of the 
base of the skull of Speotyto they remain in situ even in the dry condition. 
To return to the fronto-maxillary joint we find in all of our Vultures 
the sutural traces of about the upper third of the intermaxillary, per- 
sisting, with a line between them, indicating to us that the process of 
the bone is bifid, as we found it in the Tetraonidce. There are no good 
evidences that the ethmoid has not been completely hid beneath the 
frontals and the other segments that surround the point where it is 
sometimes superficially observed in other birds. In this position we 
find a more or less marked depression occurriug in all of the Cathartidce, 
caused \>y prominence of the iroutals behind and alike elevation of the 
culmen in front, and the lacrimals and nasals on either side. This de- 
pression is shallow and broad in Pseudogryphits and Cathartes ; deeper and 
more decided in Rarcorhamphus and Catharista, where a slight median 
elevation in the latter, better shown in G. papa than any, gives it the 
appearance of being double. From this point the upper and convex 
surface of the nasals, and the wide intermaxillary, cause the osseous 
culmen to start broad and spreading — to rapidly contract again between 
the capacious nostrils, then suddenly fall roundly convex to the tip of 
the beak, after first passing over a rise that occurs with greater or less 
abruptness just in front of the anterior margins of the peripheries of 
the nasal apertures. This is best seen in Sarcorhaniphus, and less de- 
cided in Pseudogryphus than any of the others. It is really the upper 
culmenal depression that persists down the sides of the bill to cause the 
"swell" at its extremity in the Condor and Carri n Crow. (Fig. 118, 
PI. XX.) The osseous tomia of the superior maxillary are sharp from 
a point taken below the center of the nostril, until they terminate at the 
point of the beak. This much of this margin at first presents a long 
convexity, then a corresponding concavity, to drop suddenly to the tip 
of the bill where the two meet anteriorly. A row of nutrient foramina 
are found at a greater or less distance above this margin in all the 
Cathartidw, with numerous other smaller ones scattered about above 
them, without any apparent attempt at order or regularity. Venations 
caused by the vessels running into them are permanently impressed 



742 



GEOLOGICAL SUEVEY OF THE TEERIT^RIES. 



upon the bone in many instances ; they are less perceptible in 8. cjrypMts 
than in any other member of the family. The remainder and nnde- 
scribed half of each torn mm running backwards is flat or slightly 
rounded, and includes on either side about the anterior fourth (C.aura) 
to the seventh of the maxillary (Pseudogryphits). Owing to the deep 
sides, the inferior aspect of the upper mandible is very much scooped 
out, and strongly reminds one of a well-shaped canoe; two sharp little 
ridges, one on either side, start from the tip of the beak here, and run 
backward as far as the palatine articulation, being nearly parallel with 
the tomial edges at the middle of their course. The maxillo-palatine 
fissure is wide and sub-elliptical in outline, terminating posteriorly by 
an opening in its arc that leads into the true inter-palatine cleft. (Fig. 
120, PI. XX.) 

In all of the CathartkJce we find just within the lower oorder of each 
nostril a diminutive bony shelf, formed partly by the palatine, partly 
by the lateral £>rocesses of the intermaxillary; following this along 




Cathartes aura, 

towards the tip of the beak we find it to terminate in a conical socket 
on either side. In C. atnrata they can be seen just within the anterior 
margin of each nostril, while in Pseudogryplius they are a full centimetre 
beyond. 

the bony nostrils in these birds are placed upon the sides of the supe- 
rior mandible and very nearly in their same planes. In form they 
assume more or less of an oval outline, being long and narrow in atrata, 
high and broad in the Californian Condor and the Turkey Buzzard. 

In Pseudogryplius and Sareorhamphus,\ess so in Cathartcs, their inferior 
and posterior margins blend with the transverse plate of the ethmo-tur- 



shufeldt.] OSTEOLOGY OF THE CATHALITID^E. 743 

binal coming from within-, this can hardly be said of G. atrata, as the 
latter bone in this Vulture is so far removed backwards into the rhinal 
cavity. The forms of the nostrils among the Cathartidw can be better 
studied from the various figures in the plates than from any detailed 
description we might give, however elaborate it might be. Thus far we 
have spoken of the irregular wing-like bone found in the rhinal chamber 
of any of the Cathartidw as the ethmo-turbiual, and it certainly fulfills 
the functions of that bone as found in others of the class, where it occurs. 
Not having the young of any of the Vultures at hand, however, we can- 
not say that its representative here ossifies as a separate segment, 
though to all appearances it does. 

Huxley seems to refer to it when he says : " In the genera Cathartes 
and Sarcorhamphus the cleft between the thin and scroll-like maxillo- 
palatines is very deep and wide, and the ossification of the septum is 
small in extent, and only forms a sort of bridge over the deep and wide 
valley between the maxillo palatines." — (Classification of Birds, Proc. 
Zool. Soc. Lond., 1867, pp. 441.) 

It consists essentially of a horizontal plate of bone extending across 
the nasal cavity to the rear of the nostrils, and about half way between 
the roof of the rhinal chamber and the palatine plates beneath. Mesi- 
ally it throws out a sharp x>rocess in front (C. aura), which curls up in 
Pseudogryphus (Plate XVI, fig. 106), and another plate, that reaches to 
the roof of the cavity, where it extends backwards to meet the ethmoid 
in C. aura and C. atrata, and in all of the species spreads out more or 
less laterally, thus forming a strong abutment above, while it divides 
the space into two, that by the curling edges of the plates appear like 
true nostrils within. 

The horizontal plate blends with the inferior borders of the external 
nasal orifices, and at their posterior peripheries throws up another brace, 
forming a foramen at these points in Pseudogryphus (fig. 106), and in 
the other species, but not so well marked (figs. 116, 118). On either 
side below the horizontal plate, lateral wings are developed, that curl 
inwards and backwards, reaching forward to be inserted with the pala- 
tines, showing an elliptical foramen in their sides just before arriving at 
the articulation. No doubt but this irregular ossification that we have 
been describing affords additional surface, when it is covered with the 
nasal mucus membrane, for the ramification of the nasal nerves. We 
have in our possession a specimen of a skull from Catharista at rata that 
shows extensive and distinctive necrosis of this region, the infraorbital 
bars being destroyed for their anterior thirds, with a terrible loss of 
structure to the ethmoid and lacrymals and the palatine plates beneath. 
Can we account for this state of affairs by saying that it is due to the 
entrance of putrid matters while feeding upon carrion through the 
apertures in these ethmo-turbinals? We believe that the disease in this 
Vulture before us must have eventually resulted in its death. It may be 
the cause of death among many of its kind, and, if so, would be another 
cause for the decrease in birds, and constitute an item to be taken in 
connection with Henshaw's interesting treatment of this subject. (Nutt. 
Ornith. Bull., Oct. 1881, pp. 189.) We would add that on the under side 
of the horizontal plate of the ethmo-turbinal we always find little pits 
existing, the only exception to the rule being, in the material we have — 
the cranium of Pseudogryphus. They are grouped without any particu- 
lar regard to any special arrangement, varying in specimens of the same 
species. 

The elevation existing immediately to the rear of the fronto maxillary 
articulation has already been alluded to as a feature of the superior 



744 GEOLOGICAL SURVEY OF THE TERRITORIES. 

aspect in the skulls of our Cathartidce from tliis locality as far back as 
the well-defined line limiting muscular attachment on the posterior 
aspect of the cranium and between the upper boundaries of the orbits. 
The surface of the skull presents a very evenly distributed convexity, 
it being most decided as it slopes away over the auricular entrances 
on either side. The median groove, present in so many of the class, 
is absent. The surface exhibits nianj- osseous venations, the majority 
of which run to the foramina that exist in an irregular double row, 
removed by a few millimeters from the orbital peripheries. These fora- 
mina lay along in a shallow groove in these localities. From them 
the bony and sharp-edged brows overhang the orbital cavities below 
to a greater or less degree : we say for a greater or less degree because 
this is one of the characteristics of the skulls of these birds that seem to 
vary in every individual and in every species. As a prominent instance, 
look at the two crania of Pseudvgryphus that we have before us; in one 
the superior orbital peripheries are jagged and thin, coming very close 
to the row of foramina described above, being only a little over 2 centi- 
meters apart, measuring between points in the two lines that are the 
nearest together, while in the other they are rounded and arched over 
the orbital cavities, thick and heavy, the edges being nearly 5 centi- 
meters apart, lending to the general aspect of the skull a far more 
raptorial look, as the osseous brows are thus made to be permanently 
arched and overhanging. This variance in the integrity or complete- 
ness of the vaults of the orbit may be due to the age of the specimen — 
the older the individual the more complete the roof of his orbit may 
be; but we have nothing to offer to sustain any such theory. 

Eidgway in his "Outlines of a Natural Arrangement of the Faleon- 
idce' 1 [read before the Philosophical Society of Washington, April, L875] 
gives us some very interesting and valuable studies upon the lacrymal 
audits superciliary process as found among these genera, illustrating 
his remarks with very good outline drawings of the species treated. 
"We find that the lacrymal is subject to a great many changes as to its 
form, its method of articulation, the bones it comes in contact with, and 
so on. As these changes are quite constant for the species where they 
occur, they become valuable as points of distinctive differences for 
diagnostic purposes. Among the majority of the Hawks and Eagles the 
superciliary process stands out from the head, at a varying angle for 
different species, and may have articulating with its extremity an " ac- 
cessory piece" (Harpagus bidentatus). 

In Micrastur brachypterus so long and prominent has this supercil- 
iary process become, that, aided by the intervening membrane, it makes 
up the major part of the orbital vauit. 

In Polyborus auduboni we find the body of the bone, freely articu- 
lating with the entire border of the wing of the ethmoid and three- 
fourths of the inner margin of the supercilary process, freely, though 
very intimately, meeting the frontal and nasal; the membranous inter- 
space being limited. 

The next interesting step we observe in the cranium of Gypogeranus 
serpentariiiSj where the superciliary process forms much the larger share 
of the entire bone, while the body becomes a mere inbent osseous bar 
that touches the ethmoidal wing at the angle. Here the inner border 
of the superciliary process meets the nasal and frontal segments for its 
whole length; the union being very close, but the suture plainly vis- 
ible. 

In Keopliron percnopterus the arrangement leans more towards the 
Falconidw than towards the Cathartidce^ though there is a positive step 



SHUFELDT.J 



OSTEOLOGY OF THE CATHARTID.E. 



745 



vulture-wards. In this bird the superciliary process has shrunk up in 
size, articulates principally with the margin of the nasal, while the body 
of the bone below engages the entire border of the ethmoid, all the 
natural traces being evident. The roof of the orbit is formed, by the 
frontal. A very decided change takes place among the Cathartidw ; 
here complete anchylosis with the surrounding bones has taken place, 
obliterating all traces of original individuality on the part of the lacry- 
mal. By its aid the great width is attained across the fronto-maxillary 
articulation. (PI. XXIV, fig. 127.) Its body below knits with the 
ethmoidal wing, and is produced downwards, backwards, and outwards 
so as to almost toucli the infraorbital bar, as a club -like process in Sarcor- 
hamphus gryphas and P. caiifornianus. Its outer side shows the groove 
for the lacrymal duct. By the assistance of the lateral extensions of 
the ethmoid, these bones make a very substantial partition between the 
orbital cavities and the rhinal chamber; above and between the two 
there is an opening for the passage of the nasal nerve and vessels. 

It is difficult to say, from an examination of the skulls of the adult 
birds, whether the vomer exists in them as a separate ossification or not; 
we are of the opinion, however, that it does not; at any rate, there is 
no such bone present as we find among many of the Ducks and Geese 
(it must be remembered, however, in this connection that this delicate 
little oone is often lost in the maceration and preparation of avian 
crania). » 

The ethmoid (the mesethmoid of Parker) meets the vault of the rhinal 
space above in a spreading abutment; from this point it takes a direction 




Catharista atrata. 

downwards and backwards in the mesial plane, to become consolidated 
with the extremity of the basi-presphenoid, below. Behind, by extension 
of its median osseous plate, it assists to complete the orbital septum, 
w-hile laterally it develops on either side alar productions that amalga- 
mate with the lacrymals, as pointed out above. On its anterior face we 
have presented us a sharpened edge, extending up and down the bone 
in the mesial plane. The lateral wings in G. aura are removed far to the 
back, and they have from their lower boundaries horizontal plates de- 
veloped that reach forward beyond the basi-presphenoidal tip, to the 



746 GEOLOGICAL SURVEY OF THE TERRITORIES. 

anterior base of the bone. This is a constant character and is only to be 
observed in Gatkartes a u ra. The palatines are broad plates of bone, with, 
a wide fissure separating them^ anteriorly their extremities are wedged 
into the anchylosed articulation in common with the lateral processes 
of the premaxillary and the mamillaries. Tiie posterior and upper thirds 
of their inner margins are fashioned to glide over the iuierior and 
rounded surface of the sphenoidal rostrum, the articulation being a re- 
markably free one. Their posterior ends turn outwards and accommo- 
date themselves to the entire crown of the heads of the pterygoids 
(Plate XXII, fig. 120), constituting the usual pterygopalatine articula- 
tion. Descending lamina are developed from the inner margins of 
these bones, along the posterior half of the palatine clefts in all of 
the species. There is a disposition on the part of these plates to curve 
downwards, so that their outer margins arc in a lower plane than their 
inner ones, their sur laces being smooth, and as a general thing their 
salient angles rounded. G. papa, in comparison with its size, seems to 
possess the largest and widest palatine bones of anj- of the Cathartidw, 
while C. atrata has the narrowest. 

The sutural traces of the three primary segments of the infraorbital 
bar have all disappeared, and we have remaining only the strong style 
that bounds on either side the orbital cavity below. On the inner side 
of its posterior end we observe a conical tooth-like apophysis placed at 
right angles with the continuity of the bone that tits deeply into a 
socket intended for it on the lateral aspect of the quadrate. The distal 
extremity of this malo-zygomatic link is firmly wedged into the articu- 
lation described above, being superior to the palatine, and slightly 
dilated in the horizontal plane, as the maxillary usually is. This bar 
has a gentle fall from before backwards until it arrives at the os quad- 
rature In Pseudogryphus it is much compressed from side to side, and 
presents a decided vertical swell in its anterior third. (Plate XVI, fig. 
106.) 

So large are the quadrate bones in the skull of any of these Vultures 
that they form one of the most prominent features on the lateral view 
of the cranium. The u mastoidal process" looks very much as if it 
might originally have been a broad lamina of bone, facing directly 
forwards, but subsequently had been seized by the condyle, twisted 
one-third upon itself, and its articular facet placed in the concave 
depression in the squamosal; the long diameter of this latter lies in an 
oblique direction, and if its imaginary line were produced it would pa.^s 
from the upper and outer margin of the auricular process towards the 
occipital condyle. This articular facet on the mastoid process is long 
aud narrow, being convex from side to side, as well as from above down- 
wards. The orbital process of either quadrate, though it looks very 
formidable from a lateral view, consists merely of a very thin, though 
broad, oblong lamina of bone, projecting at a right angle from the stouter 
mastoid process into the orbital space. Its inner extremity is finished 
off by a little raised rim. At its base, above the inner condyle of the 
mandibular end, we have presented to us for examination the sub-ellip- 
tical convex facette for the pterygoid, a special elevated crest being 
thrown out to support it. In all, with the exception of aura, a marked 
depression occurs just anterior to the articulation and immediately 
above the inner mandibular condyle. The Californiau Condor seems 
to have this characteristic best bhown. The condylar surface on the 
under side of the quadrate intended for the lower jaw is, as usual in so 
many birds, divided into two irregular, undulating facets, separated by 
a mid-depression ; the long diameter of the whole being situated trans- 



shufeldt.] OSTEOLOGY OF THE CATIIART1DJE. 747 

versely. Quite a decided constriction exists between the mastoid and 
orbital processes and the mandibular end. The portion bearing the 
outer condyle is produced, outwards, forwards', and upwards, as a sub- 
cylindrical, stout apophysis, having in its extremity the deep, conical 
pit let for the reception of the process upon the squamosal end of the 
infraorbital style. 

The pterygoids are horizontally compressed and exhibit upon their 
mesial edges the elongate facettes for the pterapophysial processes of 
the basi-sphenoid ; these facettes are towards the anterior and broader 
ends of the bones. The posterior extremities of the pterygoids are con- 
stricted and twisted upon themselves, so as to bring their articular fac- 
ettes to meet those that were desciibed for their reception upon the 
quadrates, while the anterior ends are dilated to afford the necessary 
articular surface for the palatines. These bones do not meet anteriorly 
in any of the Cathartidw, but form the usual palato-pterygoidal articular 
apparatus for the rostrum of the sphenoid. 

We cannot state at present with any degree of confidence whether 
the orbito-sphenoids develop in these birds from separate centers of 
ossification or not; at any rate, as a general thing, the orbital septum 
is quite complete in the smaller varieties of the family, but in the Con- 
dors and G. papa large and more or less circular deficiences, most usually 
only one, exist in this partition, about its middle. This sometimes fuses 
with the perforation that is found just anterior to the optic foramen. 
In all of our skulls of G. atrata this iuter-orbital wall is entire and quite 
thick, but in all, with the exception of this Vulture, it is in either orbit, 
produced out, by division above, in connection with the ethmoidal wings, 
to the vault, and in so doing leaves a covered duct (double in Pseudo- 
grypkus) for the passage of the first pair of nerves from the rhinence- 
phalon to the rhinal chamber beyond. So deficient are the walls of this 
osseous duct in G. atrata that it is really reduced to a canal. Even in 
Cathartes aura we occasionally find foramina, due to the too great thin- 
ning of the walls, present, and opening into the passage. In the back 
part of tlie roof of the orbit we find the usual circular foramen for the 
orbital vein, with a shallow groovelet leading from it in all of the Gath- 
artidce. 

In Neophron this arrangement is like it is in the Falconidce, i. e., an 
open channel is provided for the nasal nerves. 

The orbital septum sustains one other perforation, already referred 
to, and can be best described by saying it is a piercing of the partition 
immediately in front of the orbital foramen proper, thus allowing very 
nicely the passage of the branches of the crossed optic chiasma. Other 
nervous foramina are found to be distinct. The only example of the 
sclerotal plates that we have are a set from the eyes of a specimen of 
C. aura. In this Vulture they number fifteen in each eye, are very broad, 
overlapping each other by about one fifth of the extent ; their corneal 
margins are turned outwards, while their sclerotal ones are reflected in 
the opposite direction. We have but little doubt that when opportu- 
nity for examination offers, this description will apply very closely for 
these platelets among others of the Gathartidce. 

All of the cranial segments, or rather such of them as usually go to 
form the brain-case, have become thoroughly anchylosed in this family. 
Original sutures, boundaries, and land-marks have disappeared in the 
adult birds, leaving not a trace behind them to acquaint us as to tbe 
position of the primoidal elements. A study of the superior aspect of the 
skull has already been submitted to the reader and its leading features 
sufficiently well dwelt upon; now, viewing the cranium from a lateral 



748 GEOLOGICAL SURVEY OF THE TERRITORIES. 

standpoint (see plates of skulls), it presents for our examination, in addi- 
tion to other points that we have already considered, 1. The entrance to 
the organ of hearing; 2; Two prominent processes; 3. The general sur- 
face. 

The posterior boundary to the aural aperture is formed by a strong, 
raised, semi elliptical rim, the upper border of which is produced for- 
wards in all of the Gathartidce to terminate in a horizontal and conical 
apophysis, one of the two mentioned above. (Fig. 113, Plate XX.) This 
flaring entrance to the ear gives us a very good opportunity to examine 
into some of the osteological features of this organ as seen in the vultu- 
rine head. We observe the funnel-shaped opening of the Eustachian 
tube; the foramen leading to the inner ear, as well as several vascular 
foramina. 

In fact, in our California Condor, so shallow is this outlet that by the 
inexperienced it might be easily overlooked as not forming any part of 
the aural apparatus. 

The remaining process found on the lateral aspect of these skulls is 
above the one we have just described, and is directed almost directly 
downwards, having only a slight inclination outwards and forwards. 
We are of the opinion, and believe our reader will agree with us after 
an examination of the segmented cranium that we presented in the 
Tetraonidce, that this process was no doubt developed by the alisphe- 
noid. It becomes compressed from side to side in Neophron, still more so 
in the Secretary Vulture, while it is positively wing-like in Aquila cana- 
densis and in the majority of the Hawks and Owls. The intervening 
valley between these two processes is roundly concave, as the squamosal 
surface of this lateral aspect is generally convex outwards above and 
slightly concave below, forming, as it does, the temporal fossa on either 
side, and showing a limital muscular line above. The upjjermost of these 
two processes is called the sphenotic. 

The posterior wall of the orbital cavity is quite smooth and concave 
from above downwards in G. aura and G. atrata, less so in G. papa, while 
in Pseudogryphus and the South American Condor it is nearly flat and 
slopes away rapidly towards the sphenoidal suture, being marked by 
several transverse lines or ridges. The "foramen ovale" is unusually 
large in these birds, and is to be found rather low down in the orbit, 
almost hidden iu the shadow of the great quadrate bone on either side. 

Passing to the basi cranii we find in Gathartes and Gatharista the fora- 
men magnum to be nearly circular in outline, while in Pseudogryphus mid 
Sarcorhamphus its vertical diameter is the longer. The bony walls of the 
back of the brain-case project beyond this important aperture iu the 
median line and with a gradually lessening amount on either side, due 
to the fact that these birds all have long and markedly prominent " cere- 
bellar prominences," which rise, dome- like, above the foramen of the oc- 
ciput. This is strikingly the case in Pseudogryphus, in which, as among 
others of the Gathartidce, the inferior border of the cerebellar prominence 
and the superior arc of the foramen magnum lie in the same line, which 
line slopes away on either side to terminate in theparoccipitalsor lower 
angles of the raised ridges that bound the ears. This line or ridge forms 
a striking feature in rear views Of the skulls of the Cathartidce, and is 
present to a greater or less degree in many of the diurnal and nocturnal 
Eap tores. The condyle, situated at its usual site in the median line, fairly 
on the lower border of the foramen magnum, is broadly and transversely 
elliptical; in fact, it may be said to be semi-ellipsoidal in form andnon- 
pedunculated, though prominent. It has a faint median notch above in 
all except Pseudogryphus, where it is very feebly marked, if it occurs at 
all. We have specimens of C. atrata before us in which the occipital 



SHUFELDT.] 



OSTEOLOGY OF THE CATHAETID.E. 



749 



condyle is fully as large as in our specimen of Aquila canadensis ; indeed, 
as a rale it seems to be larger throughout the Vultures than in the 
Falconida\ 

In Polyborus tharus it is completely sessile and hemispheroidal in 
form, being still smaller, and in the Grows and Jays we know it is com- 
paratively still more diminutive. The area that includes the foramen 
magnum and the occipital condyle, and is bounded laterally by the ele- 
vated aural ridges, is depressed below the surrounding points in the 
basi-cranii of all of the Cathartidw. In this space on either side we dis- 
cover the usual venous and nervous foramina, the precoudyloid foramina, 
and those for the vagus and jugulars, the latter group occupying the 
base of a special depression for themselves. 

We find in many birds, beyond the condyle, on either side, a descend- 
ing tuberous process ; they show pretty well in a fine specimen of the 
Canada Goose that we have at hand, while they are entirely absent in 
our skull of Circus, and only moderately developed among the Eagles. 
In the Cathartidce, however, this pair of processes become the leading 
feature of the base of the cranium, though it must be remarked that they 
are not always equally well developed, for w r e have crania of C. atrata 
in our possession in which one would hardly be struck by them as 
worthy of particular notice. In one of our skulls of S. gryplms, however, 
and another of the Californian Vulture, these processes are remarkably 
well developed, being great tuberous projections that spring from ex- 
tensive bases, taking a direction downwards, outwards, and a little back- 
wards. Their inferior extremities seem to be designed for muscular and 
ligamentous attachment. In the accompanying cut, Sp designates this 
pair of processes, and Tp 
a quadrate bone. This fig- 
ure is life-size from the 
smaller skull of the two 
specimens we have of our 
Galifornian Vulture; it 
illustrates many of the 
points that we have just 
passed over in our descrip- 
tion of this view of the cra- 
nium. These basal pro- 
cesses form the outer angles 

of an isosceles triangle, the f' ^^f /jC^\ V^ "\ ?Tk\ HP /""^v 
apex of which is the lip 
beneath the Eustachian 
tubes (Plate XXII, fig. 120, 

C ailYa) and the Sides the Rear view of cranium of Pseudogryphus califomianus. 

lateral portion of the basi-crauii. The Eustachian tubes open at their 
usual site in one capacious common aperture, the gutter leading from 
the entrance of which is seen on the basi-pre-sphenoid. These tubes are 
conical, the apices being at the anterior opening; it is not uncommon in 
C. aura to find these passages not closed in by the anterior bony wall, 
and the general aperture is quite wide in Pseudogryphus. The foramina 
for the passage of the carotids are found near at hand in the recess. On 
either side of the Eustachian groove, jutting from the base of the sphe- 
noidal rostrum, we observe the pterapophysial processes ; they are 
directed forwards, outwards, and a little downwards, each being crowned 
by an elliptical facette, the major axis of which in each case is nearly in 
the horizontal plane. We have already dwelt upon the fact as to how 
these apophyses meet and articulate with the pterygoids. They are 




750 Geological suevey of the territories. 

more prominent in Gypogeranus than they are in the largest of oar Con- 
dors, while in Neophron per cnopter us the pterygoids stand well away from 
the basi-sphenoid and no such processes are present. In Polyborus not 
the slightest evidence of them exist, but in Aquila and Circus we find 
diminutive sharp spines jutting from the usual points, but they do not 
possess articular extremities, nor do they ever meet the pterygoids in 
these birds. It will be remembered that we found them in quite a num- 
ber of the American Owls, and we believe they are present in all of them. 

Gently inclined upwards, very long and robust, we fiud the basi-pre- 
sphenoid in all of the Cathartidce, carrying out its functions in the me- 
dian cranial plane. Above it has become thoroughly blended with the 
interorbital septum and the mesethmoid, below it is rounded and smooth, 
and affords the common articular surface for the pterygoids and pala- 
tines. In Pseudogryphus it has a length of four centimeters, and in C. 
aura develops a little spine anteriorly that projects beyond the meseth- 
moid. 

We show in our cut of the rear view of the cranium in Pseudogryphus 
how the region of the occiput is bounded by the superior muscular line, 
both laterally and above; this line is well marked in all of these Vul- 
tures. The lines at the sides are quite ridge-like in the California Con- 
dor, parallel and in the vertical plane, while the line forming them above 
is a long, shallow arc, with its concavity towards the cerebellar promi- 
nence; this is also the case in C. atrata. In Cathartes aura the side 
lines are curved outwards, while the superior line is broken at its middle 
point, which point is carried down on the cerebellar prominence for about 
one-third of its distance from above, in the median plane, where the ex- 
tremities of the broken line join it at a gentle curve on either side. 
This is nearly the pattern as seen in G. papa, but in 8. gryphus we again 
find it as we described it in the Northern Condor, only we have in the 
former a slight inclination for the point to come down on the/promi- 
nence. 

If we remove a section of the vault of the cranium, and this has been 
done here in G. aura and Catharista, in specimens of skulls of these birds 
that have been in collections for several years ,* we find that the internal 
and external layers or the cranial tables are very thin, and that a fair 
amount of diploic tissue is placed between them, especially towards 
the occipital region, where, as we approach the locality of the internal 
ear on either side, it becomes several millimeters thick ; the cellular 
network being more or less coarse in texture. The internal walls of the 
brain-case as thus exposed are smooth, being traversed only here and 
there by vascular tracts and grooves for the exit of certain nerve 
branches. The fossae designed for the reception of the different ceph- 
alic lobes are moderately well separated, the one that contains the 
epencephalon being the most distinct, aided as it is by the internal con- 
cavity of that external feature of the occiput that we described above 
as the cerebellar prominence ; the usual transverse groovelets do not 
mark this section here on the internal table. This distinctness is further 
assisted by thin horizontal off-shoots from the united bones of the ear- 
cell. The internal auditory foramen is unusually large and predicts a 
correspondingly good size for this important nervous branch; the same 
remark applies to the trigeminal and its orifice of exit. Eemarkab e 
depth and space is allotted to the fossa for the lodgment of the hypophy- 
sis, the "sella turcica," as this receptacle is known by in anthropomy, 

" For changes that may possibly take place, see author's remarks in Osteology of 
Lanius ludovicianus excubitorides (Bull. Geol. aud Geogr. Surv. of the Terr., Vol. VI, 
No. 2). 



snuFELDT.] OSTEOLOGY OF THE* CATHARTID^E. 751 

its posterior wall being as high as the anterior, and the cavity having a. 
depth of three or four millimeters or more. In our specimen of C. atrata, 
an elliptical perforation exists in. its hinder wall near the bottom ; the 
carotids seem to invariably pierce its base within, by two openings. 
Immediately above and anterior to it we find the optic and other 
nervous foramina spoken of when engaged with the orbital cavities. 
Passing to the rhinencepalic fossa, we are not disappointed in finding 
that this cavity is also equally spacious and well developed, and lodging 
as it does, the encephalic lobe that presides over the sense of smell, this 
fact becomes particularly interesting, insomuch that it may indicate 
extraordinary powers on the part of this faculty. The orifices of exit 
for the olfactory nerves are double in the Cathartidck, which is an excep- 
tion to the general rule. Professor Owen found the same state of affairs 
in a Vulture that he dissected, and this able anatomist made this re- 
mark upon what he observed: "In the Vulture the olfactory nerve 
is single on each side, and continued from an olfactory ganglion or 
"rhinencephalon" along the upper part of the interorbital space to be 
distributed upon an upper and middle turbinal, the latter being the 
largest."— (Anat. of Verts., Vol. II, pp. 123.) 

Along the roof of the cranial cavity, in the median line, the "longi- 
tudinal crest" is seen to pass. This may become grooved as it approaches 
its anterior termination, or for its anterior half, which indeed is the 
case in the majority of these birds; the groove dilating, and the whole 
merging into the general surface immediately before arriving at the 
conical rhinencephalic recess just referred to above. 

Fig. 115, in Plate XXI, gives us a very accurate idea of the liyoid arch, 
taken from an adult specimen of Cathartes aura. Like the other subjects 
presented the reader, it is life size; in it we notice that the ^lossohyal 
remains in cartilage during the life of this Vulture, and that the cerato- 
hyals, as two slightly curved, elliptical osseous plates imbedded in it at 
its base, articulate by the margins of their posterior arcs with facettes 
on the anterior aspect of the basihyal; they are also tangent to each 
other at their middle points in the median line. In its turn, the basi- 
hyal seems to share largely the requisite support for the broad fleshy 
tongue in this bird; to do this, it has been given an extensive horizontal 
plate -that is further strengthened by a keel below. This horizontal 
plate is concave above, dilated at its extremities, more so at the poste- 
rior one, where it produces a rounded process, mesiad, that is continuous 
with the carina beneath. This keel is deepest behind, sloping gradu- 
ally to the anterior part, where it merges into the horizontal plate and 
the thickened, elliptical facettes for the ceratohyals. Posteriorly, it is 
produced downwards as a long, slender apophysis, terminating in car- 
tilage, that no doubt represents the connate urohyal. At the sides of 
this keel, opposite its deepest part, there are, on either lateral aspect, 
cup shaped depressions to receive the expanded heads of the hypo- 
branchial elements of the thyrohyals that articulate there. These latter 
are found to be two rather robust, long bones, with upturned and cylin- 
drical shafts, that articulate in their turn with the anterior heads of the 
ceratobranchial elements behind them. The ceratobranchials are still 
more curved than their stouter companions, and gradually taper off to 
cartilage-tipped points posteriorly. 

In contradistinction to theFalconidw and the Old World Vultures, the 
members of this family are armed with much more powerful lower max- 
illae, this increased strength lies principally in the greater depth of their 
rami and consequent breadth of the symphysis, as well as the ponder- 
ous articular extremities, that these jaws possess. 



752 GEOLOGICAL SURVEY OF THE TERRITORIES. 

The vacuity, forming such a characteristic feature on the sides of the 
jaw in so many of the class, is here rarely or never present. In 
Pseud ogryphus its location is merely indicated by a shallow slit, that 
does not penetrate to the bone, though in G. papa it does for a limited 
distance along the base of a similar slit, but in our specimen of 8. 
gryphus every trace of the locality of the foramen has been obliterated ; 
again in Lathartes and Catharista narrow and faint groovelets are the 
sole indicators of its position, or the margins of the elements that origin- 
ally bounded it. 

Deep pits are found in the centers of the upper surfaces of the articu- 
lar ends; these are bounded externally by narrow, longitudinal facets, 
as do the inturned conical processes support more irregular ones. In 
the Californian Condor there is a predisposition to develop from these 
articular ends quadrate apophyses behind, in weak imitation of the 
spur-like affairs that we found so characteristic of the Tetraonichc; tbis 
effort does not seem to be so thoroughly entered into by the others. 
The under surface of either articular end is divided into two by a longi- 
tudinal ridge, continuous with the lower ramal border; of these two 
surfaces the lesser and outer faces outwards and downwards, while the 
inner and larger faces downwards and towards the median plane. 

Almost an unbroken smoothness characterizes the internal and ex- 
ternal surfaces of the sides of the jaw; this is exrended to the entire 
dentary region beyond. Even the ramal borders bounding these sur- 
faces above and below are notorious for the nice manner in which they 
are evenly rounded off, there being scarcely any evidences of the 
coranoidal projections to interrupt this general smoothness; it is only 
in the superior one, that for its anterior third on either side, and as it 
sweeps around the curve of the symphysis, that it becomes sharp, to 
correspond with the tomial edges of the mandible above. 

The depth of the symphysis in Pseudogryph us is about two centimeters, 
and the deepest part of the jaw, the ramus just beyond the articular 
ends, is LJ centimeters; for aura and atrata the measurements are equal. 
The curve that is continuous with the lower ramal borders, limiting the 
symphysis posteriorly, is parabolic in outline. Viewing the mandible 
in the Cathartidw from a lateral aspect, when it has been articulated 
with the cranium, we observe that it is bent downwards from a point a 
a little posterior to the distal end of the maxillary, from which point it 
is obliged to accommodate itself with the superior mandible. A row of 
foramina is always i^resent just within the sharp edge of the superior 
border beyond, and still within these a few others are scattered about ; 
one or two isolated, though parial, nutrient and vascular foramina are 
found at corresponding points, along the sides of the mandibles of all 
these Vultures. 

Having now passed, and we trust with sufficient thoroughness, over 
the osteology of the vulturine skull in general, and the osseous portions 
of its sense organs, may it not be in place and an advantage tons to ask 
the question here, Where have we detected any differences among these 
birds, so far as we have carried the subject ? To present, or really to re- 
capitulate, these for our reader, we will here arrange the most prominent 
departures among the species in a tabulated form, or at least such of 
them as we deem worthy of reconsideration. This method we shall 
follow, after the discussion of each of the parts or divisions of the axial 
skeleton or its appendages that may be thus separated, grouping as 
much as we can into one table, without jeopardizing its utility, defer- 
ring, however, general conclusions and comparisons for the concluding 
paragraphs of this monograph. 



6HUFELDT.J 



OSTEOLOGY OF THE CATHARTID^. 



753 



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754 GEOLOGICAL SUETEY OF THE TERRITORIES. 

Of the vertebral column. — The maimer in which the vertebral column 
of birds should be divided has been differently viewed by ornithotoinists. 
The two principal reasons for this difference of opinion, no doubt, has 
arisen from the various arrangements assumed by the free ribs at the 
anterior part of the column, and the equally diverse manners in which 
the inno uinate bones of the pelvis attach themselves to the column. 
Without entering very extensively into the literature of the iect. let 
us first examine into the question as to where the line shall he drawn 
between the cervical and dorsal vertebrae, TYe seem to have presented 
us here two very uncertain guides: the first being whether the first 
free ribs are connected with the sternum by st rnal ribs or hsemapo- 
physes. and the second upon the character of the vertebrae — that is. 
whether they have the appearance of dorsals, as we usually find them, or 
cervical as we usually recognize them. Professor Huxley sharply de- 
fines the line when he says: •• The first dorsal vertebra is defined as 
such by the union of its ribs with the sternum by means of a sternal rib: 
which not only, as in the CrocodUia, becomes articulated with the ver- 
tebral lib. but is converted into complete bone, and is connected by a true 
articulation with the margin of the sternum." (Anat. Vert. An., p. 237. - 
Professor Owen takes a different view of the subject when he states that 
••In the first and second dorsals the pleurapophysis il and 2\ terminate 
in a tree pointed end. like the -false floating ribs" of Anthropotomy; 
in the third the pleurapophysis, pZ. 3. articulates with the haemapophy- 
sis ]>. : which, in connection with its homotypes. constitutes the bone 
called •sternum."/'." (The letters given refer to a cut showiug the first 
three dorsal vertrbrae and scapular arch of a bird, in diagrammatic side 
view. (Anat. of Verts.. Vol. II. p. 15. It will be remembered that when 
we examined into the osteology of EremophUa we found sometimes that 
the second pair of free ribs, or rather freely articulated ribs, were con- 
nected with the >ternum by hiemapophyses. so that in this case some 
would claim them as true dorsals, or as dorsals any way (Owen] : while 
others could but say that the number of pairs of dorsal ribs varied. 
This state of affairs in EremophUa is no more an impossible thing or un- 
usual occurrence than the occasional presence of cervical ribs in man. 
(Owen. Anat. Verts.. Vol. II. p. 298. 

Now, among the Tciraonidrr we found another condition that proved 
equally puzzling: with them it will be recollected that there were, in 
the backbone in the dorsal region, four vertebrae that in the adult were 
completely lused together, and that the pairs of ribs that articulated 
with the anterior vertebra of this compound bone did not connect with 
the sternum by hiemapophyses. Here we must, if we consider the float- 
ing ribs in this region as cervical ribs, consider that a cervical vertebra 
has become anchylosed with three dorsals, together forming a bone that 
we believe every one would, as the writer then did. say was composed of 
dorsal vertebrae alone, in spite of the anterior ribs not joining the 
sternum by sternal ribs. One other case, and let us take a specimen of 
Asio witeonianits to illustrate it. In this bird we discover, passing from 
before backwards, that the first pair of free ribs hang from beneath the 
transverse processes of the vertebra as diminutive bonelets. as we found 
them in Speotyto. Now, the next vertebra behind this one has all the ap- 
pearances of a true dorsal vertebra, (possessing the lofty neural spine. 
etc.;: but the ribs still fail to connect with the sternum by sternal ribs. 
These three varieties may be again divided when we come to consider 
the appearance or non-appearance of uncinate processes upon these 
ribs, a condition which likewise varies. At present I do not propose to 
deliuitelv state mv views in regard to this matter, but it will be seen 



shufeldt.J OSTEOLOGY OF THE CATIIARTIDiE. 755 

from what follows that I consider the free ribs in this locality in the 
CathartidWj as well as the Falconidce referred to, as cervical ribs. This 
may be my final opinion. 

Now, so gradual is the passage from the processes on the cervical 
vertebrae from which the ribs are formed to free ribs among the Vul- 
tures, that indeed, the first cervical rib has more the appearance of a 
bony plate than a rib, yet it freely articulates with its vertebra as the 
rest of the dorsals do, and its presence makes the number of cervical 
ribs in G. atrata reckon three, Agaiu, the vertebrae possessing these 
free ribs are situated opposite the clavicles and other parts of the scap- 
ular arch, and form the " root of the neck," being so far removed from 
the dorsal region that we can only regard them as true cervical vertebrae, 
The rudimentary manner in which these ribs are first exhibited on these 
vertebrae is well shown in Plate XXI, fig. 114, cy, cy, for Cathartes aura, 
wlrile in G. atrata, as we have remarked, the par-pleurapophysial plates 
are free as ribs in the vertebrae beyond these even. 

This arrangement reduces the number of dorsal vertebrae in Sarco- 
rhampJius and Pseudogryphus to only three ; that is to say, that in these 
genera we find three lree segments in the mid-column, where the pleura- 
pophvses are connected with the sternum by the sternal ribs, while the 
vertebra immediately posterior to them is with these birds firmly anciiy- 
losed with the ossa innominata. On the same page, then, we have just 
quoted from Owen's Anatomy, it will not surprise us to find the statement 
in regard to the dorsal vertebrae that " they have not been observed to be 
fewer than four (in some Vultures) nor more than nine throughout the 
class; the latter number obtains in Apteryx, the most common num- 
bers are six and seven." Xow, if we accept this, and also this anato- 
mist's arrangement that the free ribs are dorsals and the vertebrae 
bearing them dorsal vertebrae, and there being " usually two of them," 
then it would leave us a very limited number of vertebrae between them 
and the sacrum that were connected with the sternum by sternal ribs 
(at the most two) in the Vulture referred to; so we are compelled to 
think, knowing as we do the usual condition of the ribs in these birds, 
that on this occasion the distinguished zootomist himself reckoned his 
four vertebrae as those that were articulated with the sternum by the 
intervention of sternal " haeinapophyses," and for the moment did not 
note the free ribs. This would give the same formula as G. aura and 
others. 

With the exception of the vertebrae that are grasped by the pelvic 
bones, these segments in all of the Gathartidce are freely articulated 
with each other. This is likewise the case with Gypogeranus and Neo- 
phron percnopterus, and obtains also with our Gircus hudsonius, in which 
species five vertebrae are allotted to the dorsal division of the column. 
Again, we find it in Accipiter cooperi, while in Tinnunculus sparverius and 
Polyborus tharus at least four of these vertebrae form one solid bone in 
the adult specimen ; in Micrastur brachypterus there are again five dor- 
sals, and all independent segments. 

Unfortunately, all that remains of our specimens of Pseudogryphus, 
so far as the vertebral column is concerned, are a few of the free ribs 
and several scattered vertebrae j two of these are the third and fourth 
cervical, another one from the middle of the neck j one of the last cer- 
vicals and lastly the ultimate dorsal — this latter we have devoted a fig- 
ure to, representing as it does no doubt the largest avain vertebrae of 
any living form to the northward of the range of the South American 
Condor — nevertheless, we think we may predict, almost with certainty, 
that even from these few fragmentary pieces it will be found to be the 



756 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



case, that wheD the opportunity offers for an examination of a perfect 
skeleton of this bird, that the number of segments in the spinal column 
will be the same as in Sarcorhamphus; we have been assisted in arriv- 
ing at this conclusion by a critical examination of the ribs we have, as 
well as the sternum and sacrum, that come very near to S. gryphus. 

We have carefully counted the vertebra? in the different species before 
us, and in order that the reader may have them for easy comparison, 
we present the various results in the form of a table, here subjoined, in 
which will be found other important and useful data. The number of 
sacral vertebra? must be accepted with a due amount of caution, as we 
all know it is quite difficult to decide on the number of segments de- 
voted to the "sacrum" in adult birds, after anchylosis has thoroughly 
been accomplished. 

Table for the comparison of. the vertebrce. 



Species. 



P. californimms 

S. gryphus 

G. papa 

C. aura , 

Catharista atrata 

Neophron percnopterus 
Micrastur brachypterus 



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The specimen of Mlcrastur shows an interesting phase on the part ot 
the u ha?mapophyses " of the pair of sacral ribs, for we observe the one 
on the left side articulates with the sternum by expanded extremity on 
a perfect facet as the other sternal ribs do, while on the right side the 
" hsemapophysis " of the sacral rib does not reach the sternum, but on 
the other hand lies along the posterior border of the sternal rib next 
beyond, as it does in many other birds. This gives us in this bird six 
facets on the costal border of the sternum on the left side, and only five 
on the right. 

Then, in our skeleton of Neophron we find a pair of sternal ribs, the 
last pair, springing from perfect facets on the sternum, that do not meet 
corresponding sacral ribs coming from above, but are apparently held 
in their position mainly by the intercostal muscles and membranes. 
Such facts and instances as these must make us necessarily a little 
wary, when we have only parts of skeletons in our possession, from 
which we have to decide as to the number of vertebra? or ribs m any 
particular instance. 

The Cathartidce being large-boned birds generally, we find that this 
feature is extended to the segments of the spinal column. The vertebra? 
are large, and all their various processes well marked and strong. 

In the cervical region or division of the column we find the vertebral 
canals as usual, passing from vertebra to vertebra, along on either side; 
in each segmeut the tube remains throughout more or less subcircular, 
and is closed in the ordinary manner by the parapophyses and pleura- 
pophyses of each vertebra. The protection afforded the vessels is mark- 
edly complete, for it is only in the atlas and axis that we discover slight 



shufeldt.] OSTEOLOGY OF THE CATHARTID^E. 757 

deficiencies in their lateral walls. The neural canal as it passes through 
the vertebras of the upper half of the neck is nearly cylindrical, but as 
we approach the middle of the neck it gradually becomes compressed 
from side to side, and assumes the vertical ellipse, to become circular 
again before arriving at the dorsal region. In the atlas the facet for the 
condyle of the occiput is semilunar in outline, and the neurapophyses 
are broad above, but as usual exhibits no sign of a neural spine. Below 
we commonly find a well-marked hypapophysis, though this feature is 
absent in C. atrata; laterally we have the unclosed vertebral canal of 
this bone, the processes receding from each other as sharp spiculas. 
These points are still nearer together in the axtis, and in this segment 
we find a thick, quadrate, neural spine occupying the center of the arch 
above. Below, the hypapophysis is carina-like in character, traversing 
in the median line the entire centrum of this bone. The odontoid pro- 
cess is an insignificant tip, being quite broad from side to side, while the 
postzygapophyses are tuberous lateral piojections, with the facets on 
their under aspects in the horizontal plane, looking directly downwards, 
with the anapophysial projections above, elevated int o prominent though 
blunt tuberosities. 

The facet for the third vertebra is convex from side to side, and looks 
almost directly upwards, it facing slightly backwards; the similar sur- 
face for the atlas, anteriorly, being much more extensive, twice as broad, 
continuous with the articular surface beneath the odontoid process, is 
directed forwards. Solidity and great breadth marks the third cervical 
vertebra; in it bony lr«minas connect, on either side, the pre- and postzyg- 
apophyses, an elliptical foramin being found in the surface near each 
lateral margin. 

There is a conspicuous neural spine with thickened crest, while below 
we have a quadrate hypapophysis. The vertebral canal is completely 
closed in, and parial parapophysial j)rocesses begin to make their appear- 
ance, being directed backwards ; in all of the Vultures these spine-like 
appendages are long and styliform in mid-neck, to become broad and 
tuberous as we proceed dorsalwards. 

Facets upon the pre- and postzygapophyses of this vertebras are ellip- 
tical in outline and comparatively large; the former are directed up- 
wards and a little forwards, the latter almost directly downwards. The 
anterior facet of the centrum, below and immediately outside of the 
neural canal, partakes of its usual ornithic characters; it is very narrow 
from above downwards and decidedly concave trom side to side. In 
this vertebras, the last remnants ot the carotid canal are present in all 
of the Cathartidw ; it is formed in its usual manner as we pass down the 
serial segments. In S. gryphus its first appearance is made in the eleventh 
cervical, but in the tenth in G.papa, as is also the case in C. aura and 
Catharista. More or less complete interzygapophysial bars are found 
joining the process laterally in the fourth vertebra. The hypapophysis 
of this segment is reduced to a low ridge beneath, while superiorly the 
neural spine still projects from the lamina, mesiad, as a vertical peg- 
like process. The articular facets are about as we found them in the 
preceding vertebra. 

As a rule, the hypapophysial process throughout the cervical series, 
after passing it when it is double for the carotid arteries, is found bet- 
ter marked on the next two or three ultimate vertebras. Those cervical 
vertebras that possess free ribs rarely show a distinct hypapophysis, 
but in them the centrum beneath is broad and oblong in figure, with a 
faint ridge mesiad at the usual site. 

Another suppression takes place on the part of the neural spine 



758 GEOLOGICAL SURVEY OF THE TERRITORIES. 

among the vertebrae found in mid-neck; it is but feebly developed in 
the fifth segment, still more so in the sixth, and is nearly lost in the 
few following vertebrae as we proceed down the neck. It soon reap- 
pears again, however, as a broad, knob-like apophysis, to become com 
pressed from side to side, quadrate, and finally like the anterior dorsals. 

In the Cathartidce we find a diapophysial process jutting out from the 
wall of the vertebral canal, laterally and at the anterior part of the ver- 
tebra; this character is best marked as we approach the dorsal region, 
and we find that upon those cervical vertebrae with the free ribs it is 
quite broad and exhibits a metapophysial ridge. 

Upper cervical vertebrae show long postzygapophysial processes, and 
throughout the series the arms bearing these articular facets are short- 
ened or lengthened in such a manner as to preserve the decided sig- 
moidal curve so characteristic of the vulturine neck. As we arrive at 
the middle of the cervical chain of segments, we notice that the anterior 
articular facets are barely concave, face directly inwards, and so, each 
other, occupying a position, on either side, on the bony ridge that spans 
the vertebral canal above. 

Epipleural appendages are never found upon the free ribs of the cer- 
vical vertebrae of the American Vultures, and this condition seems to 
obtain pretty generally among the Falconidw, though these ribs become 
more and more like the true dorsal ones as we advance in that direction. 

In a communication just received from Mr. Frederic A. Lucas, Ward's 
National Science Establishment, Rochester, N. Y., we learn that this 
observer found in Otogyps calvus u fifteen cervical vertebrae ; two pair 
of cervical ribs, first very small, both without uncinate processes. No 
uncinate process on last rib." This last remark probably refers to the 
sacral rib, which rarely bears an epipleural appendage. This observer 
further states that he found " a bony internasal septum (in this bird), with 
a skull proportionately shorter and higher than in Viiltur cinerea." From 
the sketch he kindly presented me the nostril has its long axis in- 
clined nearly vertical rather thau longitudinal. And, further, for Viiltur 
cinerea we find "fifteen cervical vertebrae; two cervical ribs, first smaller 
than in 0. calvus, both without processes, small uncinate process on 
last rib" (sacral?). "A bony internasal septum " was found in V. cinerea. 

Then for Gyps bengalensis, " Cervicals seventeen ; last two cervicals 
bear a rib, the first of which is 4r> millimeters long, being 40 millimeters 
longer than same rib of Y. cinerea, and 38 millimeters longer than in 0. 
calvus. Second cervical rib with long uncinate process. Last rib with 
uncinate process " (sacral ? ). "A nasal septum, but no bony projection 
within nostril, as in V. cinerea and 0. calvus. Skull much more like 
Cathartes in its shape than any other large Vulture I have examined." 
No doubt means general contour, as Cathartes, we know, has not a nasal 
septum present. 

Lateral wings are seen to project horizontally from the centrum of 
the ultimate cervical beneath, and so we pass to the first dorsal, in the 
majority of the Cathartidce, these wings still persist, but are not so far^ 
spreading, and are, as it were, drawn downwards at the expense of the 
centra, the latter becoming more compressed, the former, now attached 
by a quadrate pedicle, are true hypapophyses with flattened and ex- 
panded extremities, which latter contract, and the pedicle becomes 
longer as we approach the sacrum. These are very prettily shown in 
8. gryphus, and in all of our Vultures are a striking characteristic of the 
dorsal series. 

Rim-like projections are observed to bound the facets of articulation 
among the centra in this portion of the column; these latter are sub-corn- 



Bhufeldt] OSTEOLOGY OF THE CATHARTIDiE 759 

pressed at their middles and quite deep — this does not seem, however, 
to influence the form of the neural tube in this region to any great ex- 
tent, as it remains almost uniformly circular throughout, maikedly so 
in C. aura. Piercings into this passage ou either side for the exit of 
the dorsal nerves are made at the expense of certain parts, both before 
and behind, in each vertebra, giving rise to rather small irregular open- 
ings for this purpose. At the base of the transverse processes of the 
dorsal vertebrae, and again at their extremities, semi-circular facets 
< j xist for the capitula and tubeivula of the dorsal ribs. Those at the 
bases, are upon slightly raised elevations and look almost directly out- 
wards, those at the extremities look downwards and outwards, in aura 
and a'rata, but in S. gryplius and the California Condor almost directly 
outwards, especially in the last dorsal. 

The diapopbyses become progressively longer as we near the sacrum, 
at the same time more inclined upwards ; they are compressed from 
above downwards, being dilated at their outward extremities, where 
they bear distinet and styliform connecting metapophyses, the last pair 
being extended to the pelvis in Cathartes. 

Close and mutual locking is accomplished in this region, principally 
by a shortening of the pre- and postzygapopbyses, the facets upon the 
former facing upwards and slightly inwards, upon the latter down- 
wards and slightly outwards, so as to be nicely approximated in the 
articulated skeleton. 

Sharpened ridges beneath the transverse processes connect the facets 
for the capitula and tubercula of the ribs 5 this feature is best marked 
in C. aura and next in G. papa, less so in the others. 

Vertically elongated but shallow depressions occur above the centra 
on the anterior and posterior margins of the neural spines, for the in- 
sertion of the broad connecting ligaments; the spines themselves spring 
almost abruptly from the neurapophysial arch, are uniformly quadrate 
plates of an equal height, with thickened crests above, that become 
united at their anterior and posterior ends by a modified arrow-head 
joint, such as we described in Speotyto, where the points were more 
acute. 

Mr. Lucas tells us, in the same communication cited above, that he 
found five dorsals in the spinal column of Vultur cinerea and Gyps ben- 
galensis, six in Otogyps calvus. 

Owen clearly defines the condition of the hypapophyses in some of 
the Old World Vultures in the following words : 

From some or most of the dorsal centrums inferior processes (hypapophyses) are 
sent down, for extension and favourable origin of the flexor muscles, longi colli and recti 
antiei, of the neck. In a Vulture ( Gyps falvits) the hypapophysis is a low median ridge 
in the first and second dorsals ; to this, in the third dorsal, is added a pair of outstand- 
ing depressed plates; in the fourth the pair of plates are smaller, and, with the medial 
ridge, are supported on a common stem : in the fifth dorsal the hypapophysis is again 
reduced to a median compressed plate, but it is expanded at the eud ; the vertebra, 
which by anchylosis has become the foremost sacral, has a similar but stronger and 
slightly bifurcate hypapophysis. In both Vultures and Eagles the parial hyp.-ipophyses 
are seen to be due to modified parapophyses, which descend and are progressively lost 
in the median hypapophysis of she fourth and fifth dorsals {Harpeya, Cuv.) ; the sixth 
and seventh have only the low median ridge. (Anat. Verts, vol. II, p. 17.) 

The caudal vertebrae are very much modified, as they are as a rule, 
throughout the class ; the number possessed by each species has already 
been given in one of the tables accompanying this monograph. They 
are considered next in order afcer the dorsals, as the author believes, 
and the reader will surely agree with him, that in the present subject 



760 GEOLOGICAL SURVEY OF THE TERRITORIES. 

the sacral vertebrae bad better be taken up with the pelvic bones as an 
entirety and treated under the section devoted to the pelvis. 

In the South American Condor a complete arcade is formed by the neu- 
rapophyses of the coccygeal vertebrae, over the ultimate division of the 
myelon, and even the pygostyle is pierced for a short distance to allow 
the entrance of the nervous cord. These bony arches are surmounted by 
knob-like tubercles throughout the series, that show a very feeble dis- 
position to become bifurcated at their summits. Many of the lateral 
elements of the vertebrae are combined to form diapophysial processes, 
which, in this bird, are heavy and broad projections jutting from the 
centra on either side, bent downwards, and becoming wider and wider 
as we near the coccyx, to be suddenly suppressed in the ultimate seg- 
ment. Very faint indications of a hypapophysis occur in any of the 
first three caudals : in the fourth a cleft but sessile tubercle is seen, that 
leans forwards to rest upon the under surface of the centrum of the 
vertebrae beyond; in the last two this process becomes much larger, 
and is evidently made up of the haein apophyses of the vertebrae, for in 
each case it is pierced by a delicate haemal canal, while the true hypa- 
pophysis is still below and still exhibits the disposition to overlap the 
vertebrae beyond. The centra of the coccygeal segments of the spinal 
column in the Cathartidce, as among the class generally, are procoelian. 
In the coccygeal vertebrae of G. papa we find the same general charac- 
ters present that we have just attributed to IS. gryphus ; the principal 
differences are that the neural spines are more lofty and only the ulti- 
mate hypapophyses form a perfect haemal canal, the anterior ones being 
only grooved. Among the Vultures there is sometimes an extremely 
intimate relation existing between the first caudal vertebrae and the 
last sacral, amounting to, in some cases, positive anchylosis. 

In Cathartes aura the neural canal is complete throughout the chain 
and enters the pygostyle for some little distance: the haemal canal 
does this also below, but this latter only passes through two of the hy- 
papophyses of the last two caudals, these processes being but feebly 
developed in the others. The diapophyses in this Vulture become 
gradually broader and shorter as we leave the sacrum. Catharista ex- 
hibits about the same peculiarities with regard to its caudal vertebrae 
as we see in Cathartes aura. 

Mr. Lucas tells me that he counts seven caudal vertebrae both in 
Otogyps calvus and G. bengalensis ; he also kindly furnishes us with 
outline sketches of the pjgostyles of these two Vultures, which we give 
below as we compare them with others. 

It will be observed that, as a rule the coccyx among the Cathartidce 
is more or less parallelogramic in outline, with well-defined angles ; on 
the other hand, among the Falconidcc and the Owls and their allies 
this bony plate is drawn upwards and backwards into a rounded point; 
we give examples of this in the cuts, as well as exceptions to the rule. 

Neophron has a strong tendency Falcon-wards in this respect, less 
marked in Gypogeranus. 

We also present the reader, in each case, except in the two outlines 
from Mr. Lucas, with a posterior view of this bone, which will show the 
great variability of outline of the pygostyle, better than any descrip- 
tion could do, however elaborate it might be made. An elliptical 
foramen, in some cases two (GyjMrchus), pierce this plate at about the 
locality the last coccygeal vertebrae fused with the one next behind, 
forming, as it were, a point that the process of osseous amalgamation 
failed to fill in. This feature is rare among the Fakonida\ Among 



SHUFELDT. 



OSTEOLOGY OF THE CATHARTIDiE. 



761 



PVGOSTYLES OF VARIOUS OLD AND NEW "WORLD VULTURES AND HAWKS. 




a and a' lateral and rear view of pygostyle of Sarcorhamphus gryphus, respectively: 5 and b' the 
same for Gyparchus papa; c, c', for Cathartes aura,; d, d', Caiharista atrata ,■ e, e'\ Neophron perc- 
nopterus; /, /', Gypogeranus serpentarivs; g, Otogyps calms ; g', Vvltvr cinerea,- h. h', Tinnunculus 
sparverius; i and i', Accipiter cooperi; j, j\ Micrastur brachypterus ; k, k> Circus hudsonius; Zand 
I', Polyborus tharus. 



762 GEOLOGICAL SURVEY OF THE TERRITORIES. 

the cuts, a, a' is an outline sketch of the pygostyle from Sarcorhamphus 
gryphus ; it shows very well, and is, in fact, Typical of the parallelo- 
gramic form this bone assumes in the Gathartidce ; b and b' is the lat- 
eral and posterior view of the bone from GyparcJtus papa ; in this Vulture 
we observe that the upper portion of the bone, or rather the upper and 
posterior angle, is very much thickened. 

Gatliartes aura, shown in c and c\ still retains the usual outline of the 
other Vultures of this family, while in Catharista atrata, d and d', the 
lower and outer angle is rounded off, but the common form is still very 
decidedly retained in its upper half. In Neophron percnopterus, seen in e 
and e', we discover a marked leaning towards the form of the bone as 
it is known to us among the majority of the FaJconida\ 

Eccentricity of form stamps nearly every part of the bony frame- 
work of the African Secretary Bird, Gypogeranus, so we are not so much 
surprised to find the shape the coccygeal vomer takes on, a shape 
that we present in / and /'. That the diapophyses of the anterior 
vertebra that assisted in making this bone have been retained with 
great prominence is very evident to us, and they are shown both from 
lateral and posterior aspects. The size of the usual foramen still fur- 
ther individualizes this vertebra. 

Approximate outlines of the bone for Oiogyps calvus (g) and Yultur 
cinerea [g') are next in order given, and we note that in the first the 
tendency is decidedly to partake of Falco-form type ol the segment, 
while just the reverse is seen in the second, where it assumes a shape 
quite like the Cathartidce. 

The two more common patterns, as seen among the vast majority of 
Hawks, Owls, and Eagles, we find sharply portrayed in Circus hudsanius 
(fc, &'), and in PoJyborus tharus (?, V). On the other hand, we give nota- 
ble exceptions to this triangular style in Tinnunculus sparverius (h and 
7t'), Accipiter cooperi (i and i'), and in Micrastur brachypttrus (j and f). 
So we see that among these Vultures, as well as the Falconida?, the 
form that the pygostyle takes on can only be utilized as a distinctive 
maik in the question of differential diagnosis, when taken in connection 
with other groups of characters found either in the bird's external 
topography or its internal structure. 

The number of the dorso- vertebral ribs in any of the Cathartidce can 
easily be ascertained by consulting the table of vertebrae (the second 
accompanying this monograph), as, of course, every dorsal vertebra has 
its pair of free dorsal ribs, these being articulated in the usual manner 
with the sternum, by the intervention of the sternal ribs. The ribs as 
found among the American Vultures are very robust and strong bones, 
representing, as they do, fit accompaniments of a skeleton notorious 
for its general massiveness. As is most usual in the class Ares, the 
neck bearing the capitulum of the rib at its extremity lengthens as Ave 
pass backwards towards the pelvic end of the body, in due prox^ortion 
does the one bearing the tuberculum shorten, until in the latter we have 
the transverse process of the vertebra, in the last dorsal resting for the 
outer third of its length against the true neck of the rib apposed to it, 
and the tubercular pedicle has become sessile with the body. 

The ribs of the Gathartidce, or such of them as are found in the dorsal 
division of the column, are very broad throughout their entire lengths, 
the broadest part being found at their superior thirds; this transverse 
compression gives rise to sharp anterior and posterior borders, and long 
elliptical facets, placed longitudinally below for the sternal ribs. 

All of the dorsal ribs support epipleural appendages, in this family, 
anchylosed to the posterior margins of these bones, below the middle 



8HUFELDT.] 



OSTEOLOGY OF THE CATHARTID^. 



763 



of the shafts. We believe that this is the case in the vast majority 
of the Falconidw, including the Old World Vultures. These unciform 
offshoots are very wide-spreading and prominent, more so among the 
American Vultures than in any of the Hawks or Eagles, and as a rule 
overlap the rib immediately behind them, but never two consecutive 
ones, as in some birds. 

There are some very interesting and distinctive differences between 
the ribs of the Cathartidce and these bones, as found among the Falcon- 
idee and the vulturine Eaptores of the Continent; these differences are 
largely due to the form assumed by these very epipleural appendages, 
taken in connection with the greater breadth of the pleurapophysial 
bodies as already referred to, and shown to be characteristic of the 
family we are describing. 

W^e present afew outline, but accurate, sketches of these bones, chosen 
from several species, given in life size, as the best way to demonstrate 
these very striking and really diagnostic features. The first, marked 
m, is the rib from the anterior dorsal vertebra, taken from the left side 




of Pseudogryphus ; w, o, and p are the ribs, from the same side, of the 
second dorsal vertebrae of Catharista atrata, Neophron perenopterus, and 
Circus hudsonius, respectively. The feature which we desire to direct 
the reader's attention to particularly is the descending process of this 
pleurapophysial appendage, here best seen in G. atrata (n), though it is 
no better marked in this Vulture than in others of the family to which 
it belongs. This peculiar form of the epipleurals of the ribs in the 
Catharlidceis most decidedly marked among these bones chosen from the 
middle of the series, though it persists in a less degree in the first dorsal 
rib, as seen in m (Pseudogryphus), as well as in the appendage when it 



764 GEOLOGICAL SURVEY OF THE TERRITORIES. 

occurs on any of the sacral ribs. We know of no instance among the 
Falconidce where it ever assumes such a form; and its outline as seen 
in Neophron (o) leads us to believe that this is only an additional ex- 
ample where these birds depart from the Cathartidce and lend towards 
the Falconidce. 

The first dorsal rib in Gypogeranus serpentarius bears no epi pleural 
appendage; in the second it is broad and short, with a minute descend- 
ing process below, very close to the margin of the rib; in the third this 
unciform appendage is long and narrow, and is in contact with the rib 
for its entire length, sloping away below; in the last dorsal it appears 
only as an increased widening of the rib for a certain distance along the 
usual site of its occupancy. Sternal ribs that in descending meet the 
dorsal ribs are, like these bones, strong and robust; they possess quite 
extensive quadrate facets for sternal articulation at their lower ends, that 
when placed in situ in the skeleton cause these bones to be turned out- 
wards, so that their anterior faces are visible from a direct lateral view. 
Their posterior ends are progressively from before, backwards, curved 
in such a manner as to preserve the oval form of the chest walls, and 
are very much dilated as we proceed in that direction ; at their distal 
extremities they support the usual facets for the vertebral ribs. In the 
Secretary Vulture they become very much compressed from side to side 
as we examine them successively in the order referred to, and in this 
course, too, in some of the Falconidce, they become curved in an antero- 
posterior direction, the concave margin being in front. In these birds 
and in our specimen of Neophron the sternal ribs are seen to be much 
slenderer than corresponding bones in the Cathartidce. 

Additional protection is afforded the contents of the thoracic and ab- 
dominal cavities by ribs descending from the sacral vertebras; by free 
sternal ribs in other species, and by the meeting and articulation of both 
in others. To complete the description of these important and serial 
auxiliaries to the stability of the chest walls, we will say what we have 
observed in regard to these sacral ribs here, so that we will be left com- 
paratively free to describe that bone as a unit. 

Sarcorhamphus gryphus has three of these ribs on either side; the first 
two pair support epipleural appendages and articulate with the sternum 
through the agency of well -developed sternal ribs. The last pair are 
devoid of the unciform projections, and their sternal ribs in turn articu- 
late by their distal extremities and a small portion of their distal and 
anterior margins aloug the posterior borders of the sternal ribs in front 
of them, their points coming within about a centimeter of the costal 
border of the sternal body on either side. We believe, after an exam- 
ination of such parts of our skeleton of Fseudogryphus as refer to this 
region, that the arrangement will be found to be the same as we have 
just attributed to the South American Condor. Gyparchus possesses two 
pair of these ribs, both articulating with the sternum by sternal ribs 
that are the largest and longest of the series. The first pair have unci- 
iorm processes. Sometimes in this species an additional rudimentary 
pair are found to exist, and belong to the next vertebra beyond, but all 
the distinctive characters of the upper part of a rib have been absorbed 
by the under surface of the illiuni, so that this pair almost has the ap- 
pearance of being offshoots from the ossa innominata. 

In Cathartes aura we discover two pair, the first connecting with per- 
fect sternal ribs coming from the sternum below, and support epipleural 
appendages; the last are without them, and otherwise behave as we 
described the ultimate pair in Sarcorhamphus. Passing to Catharista, 
we find the same arrangement present as in C. aura, but in addition a 



shumsldt.] OSTEOLOGY OF THE CATHARTID^. 765 

pair of rather long styliform rudimentary ones are found, with their 
capitula, tubercula, and necks absorbed as we saw them when they 
occurred in Gyparclms, though evidently belonging originally to the 
next vertebra in order. So that among the Catliartida? the variations 
observable among the plans for the sacral ribs resolve themselves into 
the following four classes: They vary as to the number of pairs; as to 
the presence or absence of rudimentary ribs; as to the method of ar- 
ticulation of the last pair of sternal ribs, whether these descend to 
the sternum or articulate along the posterior border of the pair in ad- 
vance of them, thus constituting what might almost be termed "float- 
ing sternal ribs"; and, finally, as to the arrangement of the unciform 
processes. 

Turning to such specimens of the Vultures of the Old World as we 
have been able to learn anything about in this regard, we perceive at 
once that we have an entirely different set of "plans of arrangement" 
presented to us. In Neophron percnopterus we have a free pair of sternal 
ribs that articulate with their anterior ends on facets in the costal bor- 
ders of the sternum, just as any other pair of the series do ; no evidences 
of a corresponding pair of sacral ribs descend to meet them. 

Mr. Lucas tells me that he found in Vultur cinerea and Gyps ben- 
galensis that the ultimate pair of sternal ribs were still free as we found 
them in Neophron, but that the anterior extremities articulated with 
facets in the posterior margins of the next pair of sternal ribs beyond 
them. He further says that in Otogyps calvus "a very small floating 
rib is attached throughout its entire length to the last articulated sternal 
rib." This no doubt occurs on both sides, and is the same condition as 
we find in Gypogeranus, only the rib is longer in this latter vulturine 
Falcon. Cases of asymmetry no doubt occur among many or all of 
these various arrangements, as, for instance, in the skeleton of Micras- 
tur brachypterus that we have before us we find that on the left side six 
sternal ribs spring from the sternum, while on the right there are only 
five, the last or sixth one articulated with the posterior border of the 
sternal rib beyond it. 

Of the scapular arch, sternum, and pectoral limb. — As in the majority 
of the class Aves, the scapular arch of the Cathartidcc consists of its' 
pair of scapulas, its pair of coracoids, and its furculum, or the united 
clavicles. This group of bones enjoys the usual amount of indepen- 
dence that we find in nearly all birds, in being distinct from each 
other and from the sternum. There is a very great similarity, both 
in outline and general appearance of this arch as it is found among 
the Gathartidce, and to this we may add that when the bones forming 
it are in situ in the articulated skeleton they present a pattern that 
not only possesses a common resemblance, but is peculiar to the 
family, and differs very decidedly from the Vultures of the Old World 
and from the Falconidce. We find in our present subjects that the 
sternal extremites of the coracoids are very much expanded in a 
transverse direction, that they touch each other, mesiad, when articu- 
lated in the sternal grooves or beds designed for them. These dilated 
ends are scooped out on their posterior aspects where the pneumatic 
foramina occur, and roughened, while in front the surface is smooth, 
convex from side to side, and continuous with the general surface of the 
shaft. The inferior side is occupied for more than its inner half by the 
facet for articulation with the sternum; this is broadest mesiad, nar- 
rowing in each bone as we proceed outwards. The outer angle is trun- 
cate and presents an upturned tip of bone, and a face that is directed 
outwards. Very little sh alt can be boasted of by these bones, for no 



766 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



sooner do the fan-like lower ends commence to contract to merge into a 
shaft than dilatation immediately sets in again to form the great tuber- 
ous heads that constitute the opposite and superior extremities. More 
of a true shaft exists in Catharista than in any other of tbese Vultures, 

for the coracoids are proportionately 
longer in this species ; in ail it is more 
or less compressed from before, back- 
wards, rounded externally, sharper 
within, where in each bone it is pierced 
midway by an elliptical foramen, such 
as we found in tSpeotyto. This last 
feature is scarcely perceptible in the 
Carrion Crow. In each the facet for 
the scapula is behind and rather to- 
wards the median plane; it is placed 
transversely upon the bone, occupy- 
ing the upper surface of the scapular 
process, and is continuous with the 
shallow glenoidul facet that is seen 
on the outer aspect. The coracoids 
terminate superiorly in rounded 
heads that are flattened from side to 
side, and present upon their mesial 
aspects smooth surfaces for the broad 
clavicular limbs. The blades of the 
scapulce are short and broad, being 
curved outwards, with rounded 
points ; they never reach back nearly 
so far as the pelvis, but generally 
overlap the last pair of dorsal ribs. 
The heads of these bones are flattened 
from above downwards, curled up on 
their inner aspects, so as to afford 
surface to articulate with the points 
of the clavicular ends, while ex- 
ternally they present raised elliptical 
facets that go to complete the glenoid 
cavities of each shoulder joint. The 
entire anterior margin of a scapula 

Hight coracoid of Pseudogryphus, viewed from in is devoted tO the articular facet for 

front-, life size. the coracoid. 

The glenoid cavity formed by the approximation of these two bones 
is quite deep and extensive, and thus far we have failed to discover the 
presence of the os humero scapulare, and its assistance is apparently 
not in demand as an additional aid to retain the humeral head in the 
socket in these birds (Plate XVIII, fig. 108), the usual ligament being 
substituted for it. Our figures for the representation of the os furcato- 
rium show it to be the very type of the broad U-shaped variety or form 
(Plate XXIII, figs. 123 and 125, C. aura), and such it pre-eminently is. 
Superiorly this bone presents for examination the great flattened ends 
that articulate with the coracoids and scapulae on either side; these 
are drawn out into rounded points behind to reach the latter, while a 
limited smooth surface on the outer aspect of either limb comes in con- 
tact with a similar surface on each of the former. All of the surface 
within the U is smooth and devoid of any points of particular interest. 
Without it, and above, in the expanded heads we find the entrances to 




SHUFELDT.J 



OSTEOLOGY OF THE CATHARTID^S. 



767 



1he great air passages, for they are more than foramina, that lead into 
the bone. No hypocleidium is found, attached to the thoroughly united 
clavicles of these birds, below, but a little ridge occupies the usual site be- 
neath and a characteristic tip projects from in front in all of them. Be- 
hind the borders are rounded, in front they are sharpened and produced 
out to the point of the aforesaid tip or anterior projection. With the 
scapular apparatus in position, we find that the axis of the shafts of the 
coracoids are in line with the long axis of the sternal body ; that these 
bones diverge from each other at an angle that is equal to the angle of 
the clavicular fourchette. From behind their heads the articulated 
scapulae spring out at nearly right angles, and pass backwards parallel 
with each other, fo be deflected outwards only as we near their posterior 
points or extremities. After closing in the large u tendinal foramina' 7 
by its broad superior dilatations, the furculum dips directly backwards 
to bring its lower arch into the recess of the anterior concavity of the 
carina of the sternum, but it never touches this bone at that point, and 
its near approach seems to vary for the same species ; it is quite distant 
in the specimen of the King Vulture we present in figure 105, but in 
another it comes much nearer. 

In the vast majority of the diurnal Falconidce of this country, and, no 
doubt, in those of the Old World too, the clavicular heads have a much 
more extensive articulation with the superior ends of the coracoids than 




Circus hudsonius. 



we have just ascribed to the Cathartidce; this arrangement is closely 
followed by Neophron, and, in short, the entire scapular apparatus of 
this bird is indubitably stamped with the well-known characteristics 
that mark this arch among the Hawks and Eagles. So interesting and 



768 GEOLOGICAL SURVEY OF TIIE TERRITORIES. 

important a matter is this, that we feel sure that our reader will be glad 
to examine the cuts we present illustrating these points in representa- 
tive members of several of the families involved. The views are from 
an outer aspect of the joint in each case, showing the heads of the three 
bones of the scapular arch and the glenoid cavity. The bones are all 
lettered to correspond, and - directs attention to the furculum, to the 
scapula, and a to the coracoid. 

The figure in the upper left-hand corner is of Catharista atrata, the 
figure below is of Circus hudsonius; the remaining one is of Xeophron 
perc?iopterus. 

In Catharista we see the common plan for all of the Cathartidw, in 
which the clavicular head simply rests against the inner side of the 
coracoidal capitulum, while on the other hand, in Neophron and Circus, 
as representatives of the Falconidcc, the coracoid is actually molded to 
receive a corresponded surface on the clavicles. 

Taken in consideration with other characters, we are compelled to 
regard this arrangement of the bones of the scapular arch as still 
another valuable and reliable character differentiating these birds from 
the Old World Vultures and still further establishing Professor Huxley's 
sound classification, as far as the Cathartida -are concerned, in awarding 
them a family of their own. £To one could be more disposed to draw 
family lines in birds with greater caution than the author, nor one more 
adverse to establish such lines upon any single set of characters, chosen 
either from points in external differences or internal structure; still 
such decided distinctions as this must have their due weight, occurring 
as it does, too, in a set of bones that, taken in connection with the 
sternum, have always been regarded by ornithologists the world over 
as containing some of the most distinctive features in the avian skeleton, 
and even carried by some to such unwise extremes as to be chosen for, 
and considered of sufficient importance to even construct and base a 
system of classification upon. 

The scapular arch in the Cathartidce is far more constant in its char- 
acters than the sternum itself, a bone we will now consider; and it 
strikes us, as we glance at the many specimens before us, still more 
forcibly how totally impossible it would be to take this segment alone 
as a criterion upon which to classify the class and yet have genera and 
species of known affinities arranged anything like approaching proper 
order. My views upon the value of such single characters coincide so 
well with those of Wallace, who so long ago as 1864 appreciated and 
recognized the truth of what we have been saying, that we give some 
of his sound remarks upon this subject : 

* * * No one can be more convinced than myself of the utility of osteology, and 
especially of the sternum, in the classification of birds, and I sincerely trust that this 
great work may be brought to a couclusion (referring to a paper of M. Blanchard's). 
1 cannot, however, allow that osteological characters are an all-sufficing guide. Like 
every other character taken singly, osteology is a very uncertain and irregular test of 
affinity, and is, moreover, in almost every case accompanied by parallel external char- 
acters. Sometimes one, sometimes another part of the bird's organization has varied 
more rapidly, so that one group exhibits the most striking constancy of a part which, 
in another group, is subject to extreme modifications. The sternum is no exception 
to this rule, and by following it alone we should make the greatest errors in classifi- 
cation. For example, the sterna of the Finches and the Flycatchers are scarcely dis- 
tinguishable, notwithstanding the great dissimilarity in almost every part of the struc- 
ture of these birds — their bills, their feet, their plumage, their habits, food, and 
digestive organs. On the other hand, the sterna of the several genera of the Caprimul- 
gida? differ from each other more than do those of the most distinct families of the 
restricted Passeres. The Bee-eaters, the Barbets, and the Woodpeckers, again, are 
three very distinct families, which, in a classification founded upon all parts of a bird's 
organization, cannot be brought in close contact; and yet, their sterna, according to 



shufeldt.] OSTEOLOGY OF THE CATIJARTIDJE. 769 

M. Blaucliard, mncli resemble each other. It is evident, therefore, that the whole 
structure of a bird and its corresponding habits may be profoundly modified, and yet 
the sternum may retain a very close resemblance to a common form ; and, on the other 
hand, the sternum may undergo the important changes, while the general organiza- 
tion and. habits are but little altered. 

To prove that true affinities indicated by the sternum are also in most cases ex- 
hibited in external characters, it is only necessary to refer to the paper above quoted, 
in which the relation of the Hummers to the Swifts, and the separation of the Horn- 
bills, the Rollers, the Musophagidw, and the Parrots, from the Passeres, were pointed 
out from the consideration of such characters alone. In that paper, however, I made 
two important errors, namely, putting the Todies with the Passeres (from the de- 
scriptions given of their habits) and including the Swallows among the Swifts. The 
character of the sternum is undoubtedly of great importance in finally settling such 
points as these. 

I also at that time included the Psittaci among the Scansores ; but I am now quite 
convinced that they deserve to rank as a primary division of the class of Birds, a 
rank to which the great peculiarity of the sternum, the large brain-cavity, and highly 
organized cranium fully entitle them. 

With regard to M. Blanchard's determination of affinities from the body of the 
sternum only, without its appendages, I must remark that it often leads to erroneous 
results. For example, he says that the sterna of Merops and Tamalia do not differ 
enough to deserve a separate description, and he includes Megalosma with Tamalia iu 
one section as having the same form of sternum. He notices some differences in the 
Picidw, but remarks on their resemblance to Megalcema and to the Toucans. 

Now in all these points an examination of the entire sternum, with the furcnla, 
coracoids, and clavicles attached, leads me to very different results. * * * These 
remarks are made in no spirit of depreciation of this very interesting and valuable 
work, but for the purpose of showing that isolated characters may lead to erroneous 
conclusions from whatever part of the organism they are chosen, and that in this 
respect osteological have no positive superiority over external characters. M. Blanch- 
ard tells us 3 in the introduction to this first instalment of his work, that he proposes 
to examine successively each separate part of the bird's skeleton. His future re- 
searches may therefore seriously modify the conclusions he has hitherto arrived at. 

I cannot but think, however, that he would have produced a more satisfactory 
work if he had based it upon the comparison of the entire sternum, with its ap- 
pendages attached, and also on the cranium, these two parts being of the greatest 
importance in classification. 

It has been well observed by Professor Owen that those parts of an animal which 
have the least immediate connection with its habits and economy are exactly those 
which best exhibit deep-seated and obscure affinities. The wings, the feet, and the 
beak in birds may undergo the most extraordinary modifications in the same group 
iu accordance with differences of habits and of external conditions, while at the same 
time such apparently insignificant characters as the general coloring, the texture of 
the plumage, the scaling of the tarsi, or the color and texture of the eggs remain 
constant and reveal the true relations of the species. Thus it is that the form of the 
sternum is of such importance, since it has no immediate dependence on external 
form and habits. The Sparrow, the Flycatcher, the Wren, and the Sunbird all have 
one characteristic form of sternum, while between those of the Swallow and the 
Swift there is the greatest diversity. 

It is evident, also, that the modifications of form immediately dependent on habits 
and external conditions are generally to be seen in the skin even better than in the 
skeleton of a bird. These are principally changes of form, size, and proportion in the 
bill, feet, and the wings, which are excellent characters for distinguishing genera and 
even families ; while for determining the true affinities of isolated groups we must 
have recourse to those characters which, having no direct dependence on habits, &c , 
are often persistent in a remarkable degree. Of these, no doubt, the sternum is of the 
highest value; but there are many others of almost equal importance. * * * Now 
that true principles of classification are becoming so much better understood, we may, 
I think, hope that the chaos which has so long existed in ornithology will soon give 
way to a truly natural system which must obtain general acceptance. — (Remarks on 
the Value of Osteological Characters in the Classification of Birds, by Alfred R. Wal- 
lace, Ibis, Lond., 1864, pp. 36-42.) 

In some birds the coracoids and consequently the coracoidal grooves 
on the sternum decussate; now such a marked character as this would 
certainly have to be taken into consideration should we lay too great a 
stress upon this bone as a means of differential diagnosis in classifica- 
tion alone; and we might add to Mr. Wallace's examples such problems 
as how we would treat or dispose of Polyborus tharus and Ardea liero- 
49 H 



770 GEOLOGICAL SURVEY OF THE TERRITORIES. 

dias by the decisions of such a system, as the condition just referred to 
is found to be present in both of these birds. 

There is no distinct manubrium in the sterna of the Cailiortida\ as we 
find in Xeopliron and among the Hawks; this feature being supplanted 
by a massive and tuberous promontory in the median line, over which 
the broad concave coracoidal grooves meet at the middle point above, 
to be produced shallower, narrower, and less distinct to the deep pneu- 
matic fossae that are found on either side, just below the facet of the first 
sternal rib in the costal border, in all of these Vultures. The body of 
the sternum is oblong and deeply concave, beii g wider behind than it 
is anteriorly, and longer for its width in Catliarista than any other. 
The general internal surface is very smooth, and so evenly distributed 
is the concavity that marks it that no distinct furrow defines the po- 
sition of the carina below, as in so many other birds. All of the bor- 
ders are sharp and thin, except the anterior moieties of the lateral 
ones, that are more (Pseudogryphtis) or less (Cathartes) occupied by the 
facets for the sternal ribs. These are small parallelograms, varying in 
size according to the rib they support, placed transversely and tipped 
slightly outwards, being separated from each other by subelliptical de- 
pressions that show the pneumatic openings at their bases. Beneath, 
the body is likewise smooth, and presents for examination the prominent 
pectoral ridges on either side, originating in eminences in the middle 
of the costal borders to be produced backwards and terminate just an- 
terior to the mid-xiphoidal process, at the base of the keel. This lat- 
ter is very deep and strong in all of the Cath art idee ; commencing be- 
low and within the manubrial prominence, it is carried out, mesiad, to 
the very posterior end of the bone: its anterior margin is always thick- 
ened, but in the Condors and Gyparclw.s the entire inferior rim is fouud 
to be very much more so; in this latter Vulture, too. we observe that the 
anterior carinal margin is wider above and scooped out throughout its 
extent, having the fine median line passing down from the manubrial 
eminence to the carinal angle, that is present in all of these birds. A 
well-marked muscular line is found on either side of the keel, a few mil- 
limeters within its inferior boundary, extending from the carinal angle 
to be gradually lost before arriving at the posterior termination of this 
plate. It is less distinct in Gyparchus and the Condors. 

There seems to be no exception among the Catho.rtidce as to the fact 
that the xiphoidal end of the sternum exhibits many patterns for the 
same species; a circumstance that may be due to slight differences in 
age, but which certainly robs this bone in this family and in others 
where it may occur ot a great share of its importance as possessing 
distinctive character in this respect. It is not strange, then, that scarcely 
two drawings agree as presented us by divers authors and ornithologists. 
much less any two verbal descriptions, so that, as far as these Vultures 
are concerned, a question of accuracy has arisen on many occasions, on 
the part of one observer examining the work of another, when, perhaps, 
a mistake very rarely occurs with any of us in one description of the con 
dition in which we found the notches or foramina at the xiphoidal end 
of the sternum in any of these American vultures. 

One pattern is shown for C. aura in Plate XXIII, fig. 122. from a 
specimen secured in Wyoming by the author, and we will give a few out- 
line sketches here as the best means of showing the wonderful diversity 
that may exist, not only for the bird we have just mentioned, but for 
all, and even this characteristic is extended to the Old World Vultures 
(Xeopli ron). 



SHUFEI/DT] 



OSTEOLOGY OF THE CATHARTIDiE. 



77L 




For Cathartes aura we present the reader with the principal variations 
that we have found to exist, carefully selected from various sources and 
collections. 

Mr. T. C. Eyton, in his Osteologia Avium (London, 1867), in a half view 
of the sternum of this Vul- 
ture, found it as in spec. No. 
692, here given, only the fora- 
men was a little larger (Pate 
I, fig. 2, of Eyton's work). 
We, however, read in the text 
of this book, page 19-20 (C. 
aura), "Sternum in general 
shape similar to Sarcorham- 
pTius, but with two large fis- 
sures on the posterior margin 
next the keel, and two fissures 
exterior to them ; the remain- 
ing portions Of the Skeleton Cathartes aura, No. 689V of the Smithsonian collection. 

are very similar except in measurements." Now here is a writer that 
actually contradicts bis own drawings by the statement he makes in the 
text, and we can only believe that 
Mr. Eyton could have been led 
into such an apparent mistake by 
having several specimens of the 
sternum of this bird at his dis- 
posal, availing himself of one for 
his plate and another for his 
description, perhaps at a later 
date. 

This gentleman found his spe- 
cimen of " Cathartes niger," as we 
see from an examination of the 
plate presented, with a large cathartes aura, No. 3102 of the collection of the smith- 
elliptical foramen on either side; S0Dian institution. 
while, on the other hand, in a specimen we received from Florida, and 
shown in the cut, the arrangement is seen to be entirely different. 
Gyparchus papa apparently shares 
the same fate that C. aura does in 
this respect, for in one Ecuador 
specimen (Plate XV, fig. 105) the 
xiphoidal margin is encroached 
upon on either side of the keel by 
a broad and rather deep notch, 
each being of the same size, while, 
in a specimen of this Vulture from 
Mazatlan, this is the condition of 
the bone on the right side of the 
keel, the left having an additional 
small notch to the outer side of the 
large one. Mr. Eyton found his as A specimen of Cathartes aura at the Army Medical 
m our Ecuador bird, and describes Museum, washin-ton, d. c. 
it as having "two large open fissures," op. cit. 

Referring to these matters, Professor Owen says : 

In diurnal Raptor es the sternum is a large elongate parallelogram convex outwardly, 
both transversely and longitudinally. The manubrium is short and trihedral; the 





772 



GEOLOGICAL SUE VET OF THE TEBEITOEIE: 




lower border of the keel is convex: the front border concave : their angle r,{ anion 
rounded off. The instances where the sternum is entire have been cited: in other 

birds of prey the arrest of ossification is 
limited to very small parts of the hind 
border, usually a foramen, rarely a notch 
(.Sarc9rftantp*Ks)ott each side, one of wL: zh 
may be filled np. wholly or partially. 
Eyton figure- ~.~ -ruall notches on each 
side of the posterior border in Hierax 
bemgaleusis; and both hole and notch en 
each side in Cathartes aura. 

In our specimen of S - 

phus gryphus a faint indentation 
marks this posterior border on each 
side. bnxiu Psexdofii'i : . rstwoweB- 

: 1 notches of eqnal depth are 

OaOartaawr^JSo^^jatheSmitbaaiaaiilnsmaiafm, o v':l"~ed Oil either Side of the kc 

Eyton states tor the > r, ".it-_i AiLLt-r: 1 :/!- Condor: 

Sternum: — * * * Posterior margin in a young bird with two indentations, which 
are nearly obliterated in tbe old one. — (Ozteologia Arinm, p. 18.) 

Mr. F. A. Lucas, who kindly ex- 
amined the specimens of the 01 I 
World Vultures iu the Natural Sci- 
ence Establishment a: Rochestei 

us. writes me that. •■ You will n : 
that the right sternal foramen of 
Seophron is closed [referring t«:> a 
specimen sent to the Army Medical 
Museum): this was also the case 
w ith a second specimen, while 
third, somewhat younger, had the 
foramen open, but much smaller 

Caairista atrata.— Specimen from Florida. than the left. A Specimen of 1' 

ophron from 2s". India has both foramina open, and there are a few 
trifling differences between its skull and that of the Abyssinian speci- 
mens.'" 

The keel of the sternum in Gyp arch us has the anteroposterior curve 
alon g its lower margin, as we hud it in P- " ph us and So. rear ha mph m 
Plate XVII. rig. 1"7. P. calu nanus' : this outline is faint'y imitated 
C ~ a. but in this bird the border is not nearly so thick ev. r n in 

proportion. Catkartes has a convexity peculiarly its own. as distin- 
guishing it from others of the family. The sternum is eminently fai- 
conine among the Old World Vultures in its general form and outline. 

Professor Owen, contrasting the relative lengths :: :_r segments 
the pectoral limb as observed in the class, refers to it as found in the 
L flyers." as showing an intermediate and more harmo- 
oportion of the several segments* This is the 
?. tvith our American Vultures, for here we tind al- 
:::::: <:»f relative equipoise among arm. forearm, and 
regards lengths, but calibers of the interested loi , 




•• powerful Ptaptori 
niously balanc : 
in a marked d _ : 

most : Y":;:--t ■_■ m 
pinion, not only a: 



V 



Tc 



members of tl 
taken from Sh 
of comparison. 



;he former property in a more satisfactory manner be- 
we present a table oi the leugths of segn the 

ven in centimeters and fraoti the same, of the 

: amily under consideration, and add al mts 

rgeranus and Neophr nopterus simply for the sake 

In all-:: : $ the straght line joiniug the 



points furthest apart in distal and proximal ities was taken 



SHUKELDT.l 



OSTEOLOGY OF THE CATHARTID.E. 



773 



as the line to measure upon; in the pinion, it was the straight line let 
fall from the highest pomt in the metacarpus to the extreme tip of the 
distal phalanx, the limb being closed. 

Table shotting the length of the bones of the pectoral limb in the Cathartidce, given in centi- 
meters ; also of Neophron percnopter us and Gypogeranus. 



Species^ 



P. californianus 

S. gryphus 

G. papa 

C. aura 

O. atrata 

N. pcrcnopterus 
Cr. serperitarius . 



Humerus, 


Kadius. 


Ulna. 


27.8 


3L2 


32.8 


27. 


30.3 


31.6 


17. 


20.4 


21.6 


14.5 


16.6 


17.5 


14. 


15.1 


16. 


]4. 5 


16.7 


17.1 


18.5 


19. 


19.8 



Pinion. 



24.5 

23.5 

15.1 

14.5 

14. 

14. 

18.3 



In considering the relative position of points upon these segments 
during the course of our remarks, we must consider the bony frame- 
work of the wing as drawn up alongside the body in a state of natural 
rest, as seen in the King Vulture in Plate XV, fig. 105. The head of 
the humerus is bent not only downwards, bat anconad, the reverse being 
the case in the distal extremity of the bone ; these deflections, gentle as 
they are, and extended to a certain share of the £iiaft, give to this seg- 
ment, both from superior and lateral aspects, the usual sigmoidal cur- 
vatures. 

At the proximal extremity we find a well-developed " greater tuber- 
osity" in the form of the ordinary smooth convex and curling facet for 
the glenoidal cavity of the shoulder; below this- occurs the tuberous 
and projecting " ulnar crest n or " lesser tuberosity " overhanging a large 
subcircular fossa, at the base of which we note the many pneumatic 
perforations, to allow the entrance of air at this end of the bone. The 
radial crest occupies a position for nearly one-third (Gyparclius) or more 
(C. aura) on the superior aspect of the shaft, proximad, exhibiting all 
of its most usual points of interest. It is quite vertical, turning out- 
wards but very slightly, and strongly marked at the common sites by 
elevated muscular lines ; this crest terminates over the greater tuber- 
osity in a special broadened prominence, the continuation of its plate- 
like portion beyond. (Plate XVIII, fig. 109.) From the dilated 
humeral head, we pass to a smooth and even shaft, that presents but 
little for our examination ; it is elliptical on section throughout, the 
long or major axis being vertical, while below and nearly midway be- 
tween the extremities we observe a minute nutrient foramen that pierces 
the bone from before backwards, bearing the distal end of the hu- 
merus, the shaft gradually expands in a vertical direction, to support 
at its termination all of the characters commonly found there ; these, 
like the ones we have just left at the proximal extremity, bear out their 
ordinary ornithic types. The external condyle is raised above the bone 
as a tuberous projection for muscular insertion ; both internal and ex- 
ternal condyles are produced anconad to form outstanding and lateral 
boundaries to a shallow olecranon fossa, into which pass longitudinal 
muscular groovelets. Beyond the prominent and strongly developed 
u oblique tubercle " and " ulnar convexity," we find in all of these birds 
a triangular depression on the palmar aspect of the bone, which lodges 
pneumatic perforations already referred to. These bones are very much 
alike in their general characteristics, among these Vultures there being 
no very decided points of difference in them beyond their size ; this ap- 



774 GEOLOGICAL SURVEY OF THE TERRITORIES. 

plies pretty generally to the remaining segments of the pectoral limb. 
On the palmar aspect of the bone, at the base of the greater tuberosity, 
in Gyparchus, we find a deep pit that is not observed in the humerus 
of any other member of this family, though its site is marked in all by 
a very shallow depression. 

In the Condors we find the radius straight and nearly parallel with 
the ulna; particularly is this the case in Pseudogryphus, where for the 
distal two-thirds of its extent the interosseous space is of nearly an 
equal width; on the other hand, the bone is very much bent in G. aura, 
but heie it corresponds with a compensating curvature of its fellow, and 
little change is experienced in the interosseous space. Strong muscu- 
lar lines and decided development of the ornithic characters of the two 
extremities mark this bone. As a rule, the shaft is subtrihedral through- 
out, this being due to the prominence of the muscular lines aforesaid. 
A transverse facet occupies the entire extent of the distal aspect of its 
expanded outer end, and articulates as usual with the scapho-lunar of 
the carpus. The facet for the oblique tubercle on humerus is seen to 
be an elliptical concavity, placed vertically, with a broad, articulating 
surface, to its outer side for the ulna. Oue of the most striking features 
of the ulna that attracts our attention upon first examining this bone 
is its well-defined and double row of elevated quill-knobs for the base> 
of the quills of the secondaries; these are placed at about equal dis- 
tances apart, along the palmar aspect of the shaft for nearly its entire 
length (Plate XV, fig. 105, Gyparchus),tke fainter row being seen beneath 
them, the knobs being placed opposite each other. About its prox- 
imal end we note that the olecranon is but feebly produced, being noth- 
ing more than a general extension of the shaft, just sufficient to afford 
the necessary surface for the circular facet for the ulnar convexity on 
the humerus, the radial one being continuous with it, quadrate in out- 
line, and much shallower. Upon the inferior face of the subtrihedral or 
proximal end of the shaft, we find iu all of the Gathartidw a long ellipti- 
cal depression, that is quite characteristic, and is absent in Neophron 
and the majority of the Falconidw. Beyond this locality the shaft soon 
assumes the subcylindrical form, becomes gradually smaller in caliber 
to bear distally the trochlea surface for the remaining bonelet of the 
carpus. 

All the members of this family after they have attained their full 
growth possess but the two usual carpal segments, the scapho-lunar 
(radiale) and the cuneiform (uluare) ; these articulate with the long 
bones of the antibrachium proximad, and the trochlea surface afforded 
by os magnum, anchylosed with medius metacarpal, distad. Their gen- 
eral form varies but very little throughout the species, they bearing 
the common characteristics as we find them described for the class 
generally. 

These remarks apply with equal truth to the metacarpus (Plate XV, 
fig. 105, and Plate XIX, fig. 110 m.\ a bone that is strikingly similar 
among the Gathartidce, except in point of size, which, of course, varies 
with the species. It is constituted, as the bone usually is among birds, 
of the three united metacarpals; the short and projecting anterior one, 
or the poll x metacarpal, being the first of the bird hand ; next, the 
stout and largest of all, the second or mid-metacarpal, forming, as it 
does, the proper shaft of this compound bone; finally, the compressed 
and arched third metacarpal, thrown across, as an osseous span, 
between the proximal and distal extremities of the latter at its poste- 
rior aspect. On the palmar side of the > eeond above, Ave find a promi- 
nent projection in all of these Vultures, that in the Tetraonidm was de- 



shufeldt.] OSTEOLOGY OF THE CATHARTIDiE. - 775 

scribed as the pentosteon* that became ancliylosed at this point as the 
bird advanced towards maturity. This is a characteristic feature in 
Neophron also. The trihedral and robust pollex with its undulat- 
ing and superior facet for articulation with its metacarpal, bears below, 
in all of our American Vultures, a freely movable ungual joint; this 
appendage we have already described elsewhere, as well as the circum- 
stances attending its discovery. (Am. Stat., Nov., 1881, p. 900.) In 
tbe prepared skeleton, freed from all ligaments and soft parts, this 
joint consists simply, and in general terms lor all of the Cathariidw, 
of a compressed, usually from before, backwards, curved, and pointed 
bone, articulating upon a facette found just below the anterior margin of 
the pollex.t (Plate XV, fig. 105, Gyparelms, and Plate XIX, fig. 110 K, 
Pseudogryphus.) Now this terminal phalanx of pollex may be found 
covered by the common integuments, and not visible when the wing 
has been stripped of its feathers, e. #., in Limosa foeda and Numenius 
longirostris, or it may pierce the skin to be covered by a horny sheath, 
e. g., in many of the Ducks, Geese, and Swans. 

The particular interest which attaches to the joint in the Catliartidcvis, 
that it passes through the integument, up as far as its base, to be covered 
with a horny sheath outside, and so armed bears a very close resem- 
blance to any of the claws of the feet. This is not the case among many 
other birds, as w T e have cited above; at least the writer, who has had the 
pleasure and opportunity to examine a large number of these birds in 
company with such an astute ornithologist as Mr. Eidgway, failed to find 
it, and we are indeed moreover compelled to acknowledge here that it 

* I have proposed elsewhere the above term for this ossicle when free; I first detected 
it in the carpusof Centrocei cm; it is the fifth segment ot'the avian wrist now recognized. 
It formerly bore the name of the pisiform, which I gave it, and with which it must 
not be confused, nor with a bone so termed by Dr. Coues in his osteology of the 
Laridat. (Birds of the N. W., p. 600.) 

tThe discovery of this claw excited not a little interest at the time. Mr. Forbes, of 
London, kindly noticed it in the following letter to the editors of the American Natu- 
ralist, which contains so much of interest on this particular point, that I take the 
liberty in republishing it. 

[December 7, 1881. 
To the Editors of the American Naturalist : 

Gentlemen: I reail with much interest Dr. Shufeldt's article in your journal for 
November last, on the claw on the "index" 1 of the Cathartidce, to tbe existence of 
which he had previously called my attention when I had the pleasure of making his 
acquaintance in Washington last month. Dr. Shufeldt certainly deserves great credit 
for being the first to detect a structure, which has previously, so far as I am aware, 
escaped the notice of all observers. I may add that since my return I have been able 
to confirm the truth of Dr. Shufeldt's statements on specimens of Catharies aura and 
C. atrata in my possession. 

Allow me, as one perhaps more favorably situated than Dr. Shufeldt has been as 
regards the literature of ornithology, to call my friend's attention to Nitzsch's " Osteo- 
graphische Beitiiige zur Naturgeschichte der Vogel," published at Leipzig in 1881. In 
that 2 he will find an excellent account of the claw and phalanx in question as it exists 
in many other birds. 

Nitzsch does not seem to have observed it in the Cathartidce, but found it in Haliatiua 
albicillct. Tinnunculus alaudarius, and some others of the Falconidce. It is very conspicu- 
ous in Pandion. In fact the occurrence of such a claw is of very frequent occurrence 
in the class Aves, though by no means universal amongst them. Amongst birds in 
which it may be well seen, I may mention Struthio and Rhea, Cypselus, Caprinmlgw, 
the Pallidal and Parridw. Such a claw must not be confounded, as has been done by 
some writers, with the long "spurs" covered by epidermic tissues, formed by outgrowths 
from the metacarpal elements, of most birds, as Parra, Palamedea, Plectropierus, &c. In 
fact, the two may, as in Parra or Plectropierus, co-exist. Believe me, yours very truly, 

W. A. FORBES, 
Prosector to the Zoological Society of London. 1 

(Am. Nat., Feb'y, 1882, p. 141.) 

! The digit of the Avian manus, called "index" by Professor Owen, is now universally recognized by 
anatomists as really the pollex. 
2 Ueber das Nagelglieh der Flugel finger, besonders der Daumen," pp. 89-97. 



776 GEOLOGICAL SURVEY OF THE TERRITORIES. 

was only very recently, and when in the study of this accomplished 
naturalist, who was present, and to whom the fact was also new, that this 
characteristic occurred in FuHca, Gallinula, and no doubt in the RalUdce 
generally, not only those of our American avi-i'auna, but among their 
cousins on the other continents. Mr. James Bell, of Florida, an excel- 
lent observer of the habits of birds iu their native haunts, had that same 
morning (Dec. 28, 1S81), narrated to Mr. Ridgway how, when be was 
in Florida, he had noticed that the young of Ionomis martiniea actually 
put these claws to practical use by holding on to twigs in climbing out 
of their nests, and sometimes even suspended themselves like bats do. 

Professor Owen distinctly states that the Swan does not possess an 
external claw, when he says, after describing various claws and spurs as 
they occur among birds: '-Although the instances of these weapons, 
and the occasional use of the wings in birds not so armed, e. #., the swan, 
show them in the light of means of attack, the bones of the pectoral 
limb in birds are modified mainly for volant action." — (Anat. Verts, Vol. 
II, p. 74.) 

Mr. J. A. Jeffries does not state exactly whether in the birds he ex- 
amined the claw made its appearance through the integumeuts or not, 
in his interesting paper, upon u The Fingers of Birds." (Bull. Xutt. 
Ornit. Club, No. 1, p. 6, 18S1). This writer remarks towards the close of 
his article, that " also where there are two or three joints respectively in 
the finger there are often claws on the end, thus pointing to ungual 
phalanges." 

Since writing the article in the American Naturalist just referred to, 
I have, after more careful search among the specimens at the Smith- 
sonian Institution, found this claw in Xeophron percnopteruSj Gyps fulvus, 
and Vultur cinerea, and it. no doubt, occurs in all of the Old World 
Vultures, so that denying this fact was the most serious oversight that 
1 allowed to creep into that paper. 

So far as our knowledge carries us at the present writing, however, 
we know of no author who has directly attributed this character to the 
Gathartidw except ourselves. At the time the above paper appeared in 
the Xaturalist such able ornithologists as Dr. Cones and Mr. Eidgway 
were unacquainted with its existence, the author having the pleasure 
to call Dr. Coues's attention to the fact himself in a specimen of the 
Californian Condor; and Mr. Williarn Brewster, an ornithologist whose 
knowledge of the external characters of birds and ability the writer 
holds in the very highest esteem, writes to me as follows : 

" Your discovery of the claw on the index digit of the Caihartidce is 
most interesting and unexpected. I am no anatomist, or I might have 
anticipated you. as I have had the pleasure (?) of shooting and skinning 
both the Black Vulture and Turkey Buzzard." 

The second metacarpal supports its usual number of phalanges, the 
upper one presenting the ulnar expansion, common in so many of the 
class and here well developed, affording below a broad facet for the most 
distal phalanx of manus, the second of this metacarpal beiag very much 
like the one representing pollex, only shorter and more delicately con- 
structed. A phalanx is also freely suspended from the last metacarpal. 
This is the smallest one in the hand, being about half the length of the 
broad one of the second metacarpal, alongside of which it lies. This 
sometimes develops a tuberous process from its ulnar border, a feature 
that becomes quite prominent in Xeophron. 

Of the pelves and the loicer extremity. — We have already enumerated, 
in our table of the number of vertebra? present in the different divisions 



SI1UFELUT.] 



OSTEOLOGY OF THE CATHARTID^. 



777 



of the spinal column, the number of the sacral segments as they occur 
in these Vultures. In every instance they form a thoroughly anchylosed 
" sacrum," which in turn combines with the ossa innominata by perfect 
osseous amalgamation along the borders, more particularly anteriorly, 
for posteriorly or in the post-acetabular region the union with the ilia 
always is visible, and in some species allows a thin knife-blade to be 
inserted in the interstice by moderate 
force (Pseudogryphus). The complete 
pneuinaticityof the sacrum has already 
been cited and dwelt upon, and the 
facets for the sacral ribs, and how these 
ribs in some species unite with the ilia, 
has likewise been referred to in suffi- 
cient detail. 

Viewing this compound bone from 
above, we find that it is only in C. aura 
of the species we have before us where 
the ilia fail to meet in the median line, 
and thus conceal the common neural 
spine of the pre-sacral vertebrae (Plate 
XXI, fig. 117). In this Vulture quite 
an interspace exists between the ilia, 
8 millimeters at the narrowest place, 
which is filled in by the much com- 
pressed and very broad neural spine. 
Gyparehus makes the next nearest ap- 
proach to this condition. In all the 
others of the Cathartidw the ilia meet 
for a greater or less distance mesiad, 
and also slope downwards more rapidly 
from the line of junction, these bones 
in the pre-acetabular region of Cathar- 
tes being quite in the horizontal plane 
anteriorly. Without exception, the 
neural spine of the first sacral vertebra 
juts out beyond the anterior borders of 
the ossa innominata, as the anterior 
moiety of the segment does below. 

Cathartes aura, from the arrangement 
just described, seems to be the only one 
of the family that has a posterior open- 
ing on either side for the a ilio-neural" 

Canals, the Close approximation Of the Sacrum of Catharista atrata,- viewed from 

ilia in others of the group precluding it. above ; llfe size- 

The thoroughly united neural spines on this their superior aspect in 
this bird are also broad and flat from one extremity to the other, while 
in Catharista and Gyparchus such portions of them that show behind are 
slightly rounded from side to side, a condition well marked in the Condors. 
Keferring the reader again to Plate XXI, figure 117, we observe how 
complete is the series, on either side, of interapophysial foramina in 
the Turkey Buzzard, and how an additional row of smaller ones exist 
outside of these ; then by examining our cut of the sacrum in Catharista 
atrata (and the condition obtains next best in Gyparchus), we observe how 
it occurs among others; they are confined almost to the few ultimate 
vertebrae in Pseudogryphus and Sarcorhamphus. The anterior margins 
of the ilia are finished off above by a smooth and raised border, which 




778 GEOLOGICAL SURVEY OF TIIE TERRITORIES. 

seems to be produced over the extremities of the diapophyses of the 
first sacral vertebra? in Cathartes and the Condors. This border is car- 
ried backwards a certain distance to be lost in the true gluteal ridge, 
of which it is the anterior extension. It is less prominent in Gyparchus, 
as seen in a good life-size figure of the sternum of this Vulture given 
by Eyton in his Osteologia Avium. Posteriorly, and where the sacral 
vertebrae are visible from above, the outline of the area they appropriate, 
is lozenge-shaped, the anterior angle being in the locality of the point 
where the ilio-ueural canals terminate posteriorly in most birds; the 
posterior angle is in the last sacral vertebra, while the lateral angles are 
about opposite the ace tabula, corresponding to the longest processes 
thrown out from the vertebra below as abutments at this important 
point. The last sacral vertebra, in all of the Cathartidce, although well 
anchylosed with the one next beyond, is never completely. grasped by 
the ilia, its transverse processes always projecting a little beyond. 

The gluteal ridges meet then at a point, inesiad, in such of these Vul- 
tures as we have described the ilia meeting at a greater or less distance 
beyond the antitrochanters, tliey diverge to form bouudiug lines to the 
post-acetabular region, being carried in their course above the antitro- 
chanters, in whose neighborhood they form lateral angles to be directed 
backwards, to terminate behind in the produced processes, or process on 
either side, of the ilium. Such part of the superior pelvic surface as is 
generally known as the pre-acetabular is carried or continued backwards 
to the space between the antitrochanter and the lateral angle of the 
gluteal ridge, on either side, consequently, for description's sake, slightly 
back of the acetabulum. This well defined region is formed on each 
side by an iliac bone, and confining ourselves to one side, we find it to 
be concave from before, backwards, as well as from side to side, the 
general surface being smooth and its narrowest part just beyond the 
cotyloid ring. In the Coudors, less so in the rest of the family, the 
posterior moiety looks almost directly outwards, only slightly upwards, 
while in all the outer side of the anterior portion is directly upwards, 
and the inner nearly as in the posterior half; in Cathartes all of the 
anterior portion looks directly upwards, this Vulture having a much 
flatter and broader pelvis generally than the others of the family. 

As among Eagles and many other diurnal Raptores, the posterior 
ha^f of the pelvis of the Cathartidce, as well as the Old World Vultures, 
is bent downwards from a vertical plane passing through the bone 
tangent to the anterior arcs of the ischiatic foramina; this causes the 
post-acetabular surface or region to face backwards and upwards, it 
being bounded anteriorly by the gluteal ridges, posteriorly by a rounded 
border concave inwards, composed of the iliac margins and the last 
sacral vertebra. 

The under side of the iliac surfaces, or such portions of them as are 
seen irpon an inferior aspect, are in the horizontal plane anteriorly, 
and up to that point, as we proceed back svards to where the j deurapophyses 
and transverse processes of the first four or five vertebrae are thrown 
out to abut against and anchylose with them. The anterior face of the 
first sacral vertebra possesses all of the necessary elements to articulate 
with the last lumbar; its neural spine is quadrate and produced beyond 
the ossa innominata ; the prezygapophyses look upwards and inwards; 
the entrance to the neural canal is elliptical with the major axis verti- 
cal. In all of the Cathartidce the first sacral vertebra supports more or 
less of a well developed hypapophysis ; this process is very large in our 
specimen of Pseudogryphns, possessing lateral wings and upon its anterior 
face bearing a small facet for articulation, with a similar process upon the 
last dorsal. This plate-like hypapophysis, merging with a more feebly 



shufeldt.] OSTEOLOGY OF THE CATHARTID^E. 779 

developed one on the vertebra next behind, in this Condor, added to the 
marked lateral compression of the centra of the first three or four sacral 
vertebrae, give this bone an extremely odd appearance for its under and 
anterior half, the bone dipping far down into the abdominal cavity, a 
characteristic that is more or less evident among others of the family, 
but we do not find it so iu Neophron, an Old World Vulture that pos- 
sesses a pelvis very much nearer the Falconidce. Branches of the sacral 
nerves, the motor and sensitive roots, make their exit from the sides of 
several of the middle segments, the foramina being placed one above 
another, the largest apertures occurring in the mid-series, becoming- 
smaller as we advance anteriorly or posteriorly. 

The margins of the anterior, and at the same time horizontal, portion 
of the iliac bones are quite sharp. They converge to points at a greater 
or less distance beyond the acetabula, depending upon the species, 
where tbey suddenly become robust and rounded and continuous with 
a similar surface belonging to the u heads" of the pubic elements, or 
that portion of these bones on either side which complete the cotyloid 
rings. Opposite this, the constricted portion of the pelvis, or rather its 
narrowest part, we observe the greatest amount of enlargement of the 
neural canal to accommodate the ventricular dilatation of the myelon; 
here, too, short parapophy sial braces come from the vertebrae, to be directed 
upwards, the anterior one abutting against the ilium on either side, the 
posterior rarely, if ever, meeting these bones. 

Viewing the under side of the bone, we note that it is just beyond this 
locality that we drop into what really may be designated as the " basin 
of the avian pelvis," and here usually the next three vertebrae throw out 
only their superior processes as braces to the ossa innominata, while the 
last five (Pseud ogryphus, Cathartes, Catharista) or six (Gyparchas) sacrals 
have strong parapophyses that unite at their outer extremities with each 
other, and with the other processes coming from the vertebrae all along 
the iliac borders on either side ; of these, the first pair is the longest and 
are the ones opposite the cotyloid rings, the last two having all of their 
processes run more or less together, forming two pairs of strong rounded 
braces that are extended nearly horizontally to the " side bones." This 
pelvic basin is very commodious and deep, more particularly in the Con- 
dors and the Carrion Crow ; this condition is much enhanced by a sort 
of reduplication that takes place from the posterior and united portions 
of the ilia and ischia, forming a concave recess on either side just within 
the ischiadic foramen. 

Upon a lateral view of the pelvis, we find the acetabular ring nearly 
circular, the peripheries of the inner and outer boundaries coming nearest 
together in their upper and anterior arcs, while at their posterior and 
upper arcs they form the outline of an extensive antitrochanter, whose 
surface is directed forwards, downwards, and outwards. The greatest 
amount of surface for the articulation of the femoral head, between the 
internal and external ring, is found anteriorly and below. A stout os- 
seous pillar separates the cotyloid ring from the much larger and sub- 
elliptical ischiadic vacuity, which is posterior to it. Below and between 
the two we find the long, oval obdurator foramen, its major axis nearly 
parallel with the pubic bone, and a deficiency occurring at its posterior 
arc, where this latter element fails to meet the ischium. The separating 
and outlying bone about these lateral openings in the pelvis of the 
Cathartidce is thick and strong, more particularly about the acetabular 
ring, affording ample support for the powerful pelvic limb of these birds. 
The pubic style, after passing the obdurator foramen, is a moderately 
wide strip of bone, compressed from side to side, nearly or quite touch- 



780 



GEOLOGICAL SURVEY OF THE TERRITORIES. 




SHUFELTTT.] 



OSTEOLOGY OF THE CATIIARTIDiE. 



781 



ing for its entire length the lower ischial border, except in Gyparchus, 
where quite an interspace seems to exist. Its outer or posterior ex- 
tremity is produced well beyond the other pelvic bones, to curve inwards 
towards its fellow of the opposite side, from which it is separated by a 
varying space two and a half centimeters in our specimen of Sarcor- 
hamphus, nearly one and a half in C. aura and Catharista. That portion 
of the outer aud lateral surface of the ilium that is posterior to the 
ischiadic foramen, and below the continuation of the gleuteal ridge, 
looks downwards and outwards ; opposed to it, below, is the ischial sur- 
face looking upwards and outwards; these bones thus form a longi- 
tudinal aud shallow furrow between them, the anterior extremity being 
in the posterior arc of the ischiadic foramen, the posterior extremity 
terminating in the apex of a notch that is found between the ilium and 
ischium in the posterior pelvic margin. This notch is acute or angular 
in the Condors and the King Vulture, but rounded in the Turkey Buz- 
zard and the Carrion Crow; it is very distinctive of the Cathartidce, 
none of the Old World Vultures or the Falconidw apparently possessing 
it, it being absent in all of the representatives of these latter birds that 
we have before us, the nearest approach to it being in Gypogeranus. 

In reviewing the forms of the pelves of these Vultures and comparing 
them with each other we find, of course, for size, that the bone is largest 
in Pseudogryphus and Sarcorhamphus gryphus, and indeed in these two 
birds the pelves are very like each other. In general outline, and dif- 
fering from the Condors, we find the pelves of Catharista and Gyparchus 
to be nearest alike, and to this couplet the pelvis of Neophron percnop- 
terus approaches quite near, while Cathartes aura has a pelvis ot a form 
differing from all the rest, and peculiarly its own, being at once recog- 
nized by the separation of the ilia anteriorly by the broad neural spine, 
and by its greater width as compared with its depth and length. 

As already hinted, the pelvic limb of these Vultures is a very well 
developed one, and in every respect equally so with the pectoral, 
extremity, which we have described above. 

Among the species there are but few and trifling differences, except 
in point of size, though we note some departures from the typical form 
assumed by the Cathartidw in Neophron, a Vulture that has a lower 
extremity more like the true falconine birds, as it did an upper one. 

We find the pelvic limb in the Cathartidw to consist of the usual 
number of bones for birds throughout the class, and, as a matter of some 
interest and no little importance, we insert here a table showing the 
comparative lengths of these segments, given in centimeters and their 
fractions, just as we did for the pectoral limb, choosing the same species 
and specimens to measure from, so that the two tables can be compared 
and the balance between the two limbs of any species taken into con- 
sideration; this will show at once the well-known discrepancy existing 
in Gypogeranus. 



Table showing the lengths of the hones of the pelvic limb in the Cathartidw, given in centi- 
meters; also of Neophron percnopterus and Gypogeranus. 



Species. 



P. calif ornianus . 

S.gryphus 

G. papa 

C. aura 

Catharista atrata 
N. percnopterus . 
G. serpentarius . . 



Femur. 


Tibia. 


13.6 


22. 


14.8 


23. 


10.1 


17. 


7. 


11.9 


9.3 


14.8 


7.3 


11.8 


11. 


30.1 



Tarso-met. 
tarsus. 



12. 

12.5 
9.2 
6.4 
8.6 
7.5 

29. L 



782 



GEOLOGICAL SURVEY OF THE TERRITORIES. 



These measurements show us that among the Cathartidce, the Condor 
of our western country possesses the greatest extent of wing, although 
the South American one (Sarcorliamplius) has the longest legs; the same 
condition is also seen to exist between Cathartes aura and Catharista 
atrata. Also, in comparing C. aura with N.percnopterwt, we observe that 

although the measurements of the seg- 
ments of the pectoral limb are very nearly 
alike, the latter bird has a longer tarso- 
metatarsus in comparison, even where the 
femur and tibia are nearly as in the first; 
here again we find in this Old World 
Vulture a balance among the segments 
of this extremity that similates the Fal- 
conidce. 

The femur is so bent that a longitudinal 
line drawn along its anterior surface is 
convex outwards, the greatest curvature 
existing at the junction of middle and 
lower thirds of shaft. A similar line 
drawn down its inner aspect is found to 
be concave. 

The upper surface of the semi-globular 
head is in the same horizontal plane with 
the extensive articular facet at the sum- 
mit of the bone for the antitrochanter of 
the pelvis, while above this rises the 
broad and prominent ridge of the great 
trochanter: below and to the outer side of 
which we find the pneumatic foramen (C. 
aura) or foramina (Pseitdogryphus), for 
generally the species show more than one. 
The femoral head is eminently sessile 
with the shaft, and presents for examina- 
tion above an extensive though single 
excavation for the lig amentum teres. Be- 
low the characters we have just enumer- 
ated as pertaining to the proximal ex- 
tremity of the bone, the shaft rapidly be- 
comes subcylindrical, to dilate trans- 
versely at its distal or condylar end in the 
usual way. Near the middle of the shaft 
behind we observe the medullary orifice, 
and the ordinary muscular lines are tol- 
erably well produced. 

The rotular channel on the anterior 
aspect is moderately deep, rather wide, 
and of nearly the same width throughout ; 
gryphus caiij vrnianus ; life-size. it passes beneath into a shallow, inter- 
condyloid notch. In the popliteal depression, above the condyles behind, 
a deep pit exists; a few foramina are found at the bottom of it in Cathar- 
ista atrata that may be pneumatic. The fibular cleft at the back of the 
external condyle is very decided, the inner half formed by it being pro- 
duced well backwards in all of the Cathartida?, as a prominent process to 
be applied to the internal aspect of the fibula in the articulated limb. 
Slight depressions are found, one on either side, in the broad lateral 
surfaces of the condyles, intended for ligamentous insertion. If a plane 




Anterior view of right femur from Pseudo- 



shufeldt.] OSTEOLOGY OF THE CATHARTID^E. 783 

is brought tangent to the lowest point in either condyle, so as to include 
them both, it will be seen that the axis of the shaft is nearly perpen- 
dicular to such a surface, showing that the outer condylar tuberosity to 
be but very little lower than the inner, not nearly so much so as in some 
birds. 

The patella in these Vultures is of fair size, and bears out its most 
common ornithic characters, being more or less flat superiorly, convex 
in front, and divided into two unequal faces behind, the inner being 
tlfe larger; it is situated well above the cnemial crest of tibia in the 
tendon of the quadriceps fern oris. 

Quite an intimate union exists between tibia and fibula, although pos- 
itive anchylosis never takes place, along the produced and prominent 
fibular ridge on the outer aspect of the shaft of the tibia, occupying a 
good share of the upper third of this bone. Above this point the fibula 
is very much enlarged, and drawn backwards into a laterally compressed 
tuberous head, with a smooth, nearly horizontal facet above, that in none 
of the species rises much above the summit of the articular surface of 
its companion. Below the fibular ridge this bone dwindles to its usual 
styliform dimensions, being compressed from before, backwards, and 
running well down the tibial shaft into its lower third, to terminate 
in a free x>ointed end, in all of the Gathartidw, except Gathartes aura, 
though the union is very intimate at the lower extremity in the skeleton 
of Sarcorhamphus. The tibia has a large cuboid head, but the undulat- 
ing articular surface at its summit is not profoundly impressed by con- 
dylar depressions, for the trochlea? of the femur, and, indeed, the pro- 
ami ecto -cnemial ridges are but feebly developed ; the latter is produced 
fibularwards as a strong though blunt tuberosity, shielding the superior 
tibio-fibular articulation in front. The cnemial crest above these processes 
is likewise low and not raised to any extent above the general articular 
surface to which it forms the anterior boundary. 

A section of the tibial shaft, made anywhere between the distal ex- 
tremity and the fibular ridge, shows it to be broadly elliptical, and the 
entire shaft is bent so as to be convex anteriorly, concave throughout 
its length posteriorly; it expands transversely as it approaches the dis- 
tal extremity, where we find the usual points for examination found in 
the vast majority of the class. A broad and strong osseous bridge is 
thrown obliquely across the groove that is the continuation upwards of 
the intercondyloid notch, to retain the extensor tendons. The trochlea? 
are reniform in outline, placed in anteroposterior and nearly parallel 
planes, the fibular one being the broadest anteriorly; the notch sepa- 
rating them is deepest just below the bony. bridge for the extensors in 
front, while behind it is not carried very far up the shaft and becomes 
very shallow, the trochlear apparently running into one common surface. 

The bones of the leg of Neophron percnopterus are very similar to those 
found in the Gathartidw ; the principal differences seem to be that the 
pro- and ecto-cnemial ridges at the proximal extremity and the trochlea? 
at the distal are placed rather farther apart ; the bony span to hold the 
extensor tendons is the same. We mention this fact because in some of 
our American Hawks (Tinnunculus, Polyborus) it is found to be double, 
i. e., the bridge above is thrown across a wider tendinal groove in these 
birds, and from the lower margin of the span another bony piece is 
joined that is carried down to the in ter-condyloid notch. This arrange- 
ment gives one opening above and two below, one on either side of the 
last bony span mentioned. 

Unfortunately we will not be able to enter into the subject of the tarsal 
segments, and their mode of union to the bone we are now about to 



784 GEOLOGICAL SURVEY OF THE TERRITORIES 

describe, and to the distal end of the tibia, as this requires the skeletons 
of the very young of the subjects in hand, material that I unfortunately 
do not possess. 

The study of the forms assumed by the process or processes at the 
proximal and posterior aspect of the tar so -metatarsus is extremely inter- 
esting, and as important as many of the results are, the author is obliged 
to confine himself here to a hasty sketch of the general appearance of 
this tuberosity as it is found in the Cathartidm and some of the Old World 
Vultures and the Falconidce. Among all of the American Vultures it is 
a broad cuboidal process, placed at the extremity of the posterior aspect 
of the shaft, mesiad. It has leading away from it below a raised crest, 
that soon merges into the shaft, or the amalgamated mid -metatarsal. 

In Cathartes this process is sharply grooved in a vertical direction be- 
hind. This is also the case in Catharista; the King Vulture has the 
process broader transversely, the grooving shallower, with its outer and 
posterior margins slightly produced. This condition is still farther 
advanced in the Condors, while upon turning to Neophron we observe 
that it has been carried still farther, so much so that the mid-vertical 
groove is now abroad concavity and the lateral productions appear as 
separate and rounded processes. 

In Polyborus thams it is a distinct, broad quadrate plate at right angles 
to the shaft. In Tinnunculus, this plate is not distinct, or rather it is 
carried far down the shaft to merge into it in the lower third; an in- 
termediate form between the last two is presented by Micrastur brachyp- 
terus. 

Gypogeranus presents it very much the same as in Cathartes aura, 
only rather longer for its width, which is quite natural in this bird of 
a stilt-like tarso-metatarsus. An instance of its being apparently 
double is seen in Buteo cooperi, one stout and quadrate lamelliform 
process, crowned by a transverse, subelliptical plate, standing promi- 
nently out from the fibular side of the bone, while absolutely separated 
from it on the tibial side we find a smaller though eminently distinct 
apophysis with its summit slightly bent outwards, the intervening sur- 
face being broad and only moderately concave between the two. Other 
Hawks also assume this form, as Circus. A tough piece of cartilage is 
placed over this process in the Cathartidce, through which many of the 
llexor tendons pass. The summit of the bone presents two lateral con- 
cavities with a median anterior rounded tip, all for the accommodation, 
in the articulated skeleton, of the trochlear of the tibia. 

Horizontal sections made at almost auy point of the shaft are more 
or less parallelogramic in outline, and this portion of the bone is mark- 
edly straight in all of these Vultures, for we know that in many of the 
Falconidce, and the condition is slightly observable in Neophron, that 
the tarso metatarsus is often more or less bent in the reverse direction of 
the tibia above. The shaft of this, the last long segment of the hinder 
limb, is very much scooped out on its anterior and upper surface; this 
disappears as we approach the distal end of the bone, where we find the 
large foramen for the anterior tibial artery occupying its usual site. Be- 
hind, the shaft is doubly grooved, in a longitudinal direction, markings 
that are about equally distinct throughout their course, each groove pass- 
ing down alongside the plate that was described as coming from the pro- 
cess at the upper and hind end of the bone. Two foramina pierce the 
bone at its upper part, appearing posteriorly on either side of the plate 
just mentioned. The three trochlear projections that terminate this 
bone distally are large and well separated from each other, the mid 
one being the largest, standing out in front of the others, and possesses 



6HUFKLBx.i| OSTEOLOGY OF THE CATHARTIDiE. 785 

a very decided median groove that passes clear round its entire sur- 
face; this feature is usually absent on the lateral processes, of which 
the outer is the smaller; these are placed slightly to the rear of the 
middle one, particularly in the Condors, least of all in Catharista, in 
which Vulture all three are nearly in the same transverse plane. The 
concave facet for the os metatarsale accessorium is more than usually 
distinct, and this bone in the recent skeleton is attached after the com- 
mon rule by ligament merely; it is twisted upon itself, rather long, but 
not so long as in Neophron in proportion, and supports its ordinary 
toe, of a joint or phalanx and an osseous claw. 

Perhaps there is no better way of calling the reader's attention to the 
points of interest that are to be found in the feet of these birds than by 
comparing such a Vulture as Gyparchus papa, that has represented in 
these parts all of the characters of the Cathartidw, with Neophron per- 
cnopterus, that as far as we know possesses in its foot all of the striking 
characteristics of the vulturine birds of the Old World. The joints of 
the toes follow the usual avian rule of 2, 3, 4, and 5 segments to the 
first, second, third, and fourth toes, respectively. In the first or hind 
toe of Gyparchus, and in all of the Cathartidw, the proximal joint is long 
and about equally dilated at either extremity, while in Neophron the 
end that articulates by its concave trochlear surface with the os meta- 
tarsale accessorium is very much expanded transversely, while at the same 
time it is compressed from above downwards. The bony tubercle found 
at the under side of the proximal extremity of all of the claws is quite an 
insignificant affair in our Vultures as compared with the protuberance we 
find in Neophron, and, moreover, the claws are very much more curved in 
this latter bird than they are in the Cathartidw. The proximal joint of 
the inside toe of Gyparchus is long, having all the characteristics of the 
other long segments of the foot, while in Neophron it is a markedly short 
and irregular bone, having, to be sure, its ordinary articular surfaces, 
one at either extremity. This difference can be made more evident by 
simple measurement; the first and second joints of the inside toe of 
Gyparchus measure respectively 2.2 and 2.5 centimeters in Neophron 
percnopterus the same segments .7 and 2.4 centimeters, respectively. It 
is very interesting for us to know that in this matter of the shortening 
of the first joint of the inside toe Neophron follows all of the Falconidce 
or their American representatives that we have been able to examine. 
Differences in the hind toe are not so striking, the segments in both 
birds being long and proportionately balanced, but in the outside toe 
again we discover a leaning on the part of Neophron towards the Fal- 
conidce, while Gyparchus, in common with the rest of its well-marked 
family, still adheres to a proportionate equipoise in length of the inter- 
nodes, this time it occurs in the second and third joints of the toe in 
question. These we will also compare by measurement: in Gyparchus, 
first, second, third, and fourth segments measure 1.8, 1.4, 1.1, and 1.6 
centimeters, respectively; in Neophron the same segments measure, in 
the same order, 1.2, .5, .4, and 1.5 centimeters. 

In closing this monograph the writer did intend to give a general 
synoptical table, but upon second consideration believes the various 
tables already presented will sufficiently illustrate and compare, not 
only the decided generic differences* among the Cathartidw themselves, 
both as to external characters and the more deep-seated internal and 
osseous distinctions, but also serve sufficiently to individualize the group 
as a family ; so that, upon reviewing all that we have endeavored to pre- 
sent upon the osteology of these birds,, we firmly believe that the 
reader will agree with us that our investigations have not only confirmed 
the fact that the Cathartidw axe widely distinct from the Old World 



786 GEOLOGICAL SURVEY OF THE TERRITORIES. 

Vultures but enough so among themselves as to certainly warrant the 
family divisions into genera that we have followed in this paper. We 
farther believe that the time is not far distant when the fact will be 
generally acknowledged, and Professor Huxley's admirable arrange- 
ment is this regard followed, that the Otld World Yultures are only en- 
titled to the rank of a sub-family under the Falconidw. It is hard to 
perceive, even, how Mr. Sharpe can still adhere to his classification as 
given in the catalogue, where both New and Old World Yultures have 
been placed in the same family, Vidturidw, and then divided into the two 
sub-families, I. Vulturince (Old World Vultures), and II. Sarcorhampliinw 
(New World Yultures.) (Cat. Birds of Brit. Mus., Yol. 1, 1874.) Let us 
take the very good example of Gyparchus papa from the second of these 
groups, and Neophron percnopients from the first , snow-white skeletons 
of both of these birds, mounted upon their museum perches, are now 
standing before me, silent attestors,, as far as their osteology goes, of 
the violence perpetrated by such an arrangement. 

in Gyparchus papa we have the pterapophysial processes of thebasi- 
sphenoid, the peculiar arrangement of the lacrymals,.and the absence of 
the nasal septum, cranial distinctions of great importance and weighty 
while in Neophron a complete septum narum exists,, and the ptera- 
pophysial processes are missing, characters it has. in common with the 
Falconidw. 

Then, again, the differences in the scapular arch and sternum, the 
pneumaeity of the skeleton on the part of the King Yulture, a condition 
not enjoyed to any such extent by NeopHiron, the arrangement of the 
sternal ribs, and, in short, down to the very toes, these two birds are 
stamped with characters that compel us to acknowledge that they belong 
to two very different families, and to such, along with others of their 
kind that possess similar and such undoubted differences in structure^ 
they must eventually be assigned by universal consent. 

Washington^ Augmt 31* 1882.. 



788 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XT, 
FiO. 105. — Skeleton of Gyparchus papa > J, one-third the- size- ofT life. 



790 GEOLOGICAL SURVEY OF THE TERRITORIES, 



PLATE XVI. 

Fig. 106. — Skull of Pdeu&ogryplms calif or nianus, left lateral view, life size. 



792 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XTII. 

Fig. 107. — Right lateral view of sternum of P. ealifornmnu8 r from same specimen 
as iigure 106 ; size of life. 



794 GEOLOGICAL SURVEY OF THE TERRITORIES, 



PLATE XVIII. 

Fig. 10&, — Right coracoid and scapula of P. calif or nianus, size of life ; the dried 
tissues, stretching across the glenoid cavity, assist in retaining these bones in their 
proper position. 

Fig. 109, — Right humerus, anconal aspect, from the same individual ; life size. The 
dark pit at its lower side shows the entrance to the largest of the pneumatic fossae. 



796 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XIX 

Fig. 110. — Outline sketch of the left metacarpus, P. californlanus (same bird), with 
pollex (K), and the bony core of the claw it bears, in situ; life size, m, metacarpus; 
d, pollex. 

Fig. 111. — Furculum of Pseudogryphas californlanus, size of life, from the same spe- 
cimen ; viewed from the left side, showing the positions of the pneumatic foramina and 
the roughened surfaces for the attachment of ligaments. 

Fig. 112. — The last dorsal vertebra, from the same bird, life size, posterior aspect, 
showing the facets that articulate with the first sacral vertebra. pf } large pneumatic 
foramen. 



79 S GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XX. 
Tig. 113. — Left lateral view of life-size skull of Sarcorliamplius gryphus. 



800 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XXI. 

Fig. Il4. — The three last cervical vertebrae from Cathartes aura, seen from below, 
showing the formation of the cervical pleurapophyses from the processes of these 
vertebrae by gradual metamorphosis as the bird passes from young to maturity ; speci- 
men size of life, cy, ribs that remain free in the adult and after the change just re- 
ferred to is complete. 

Fig. 115. — Hyoid arch of the same bird. 

Fig. 116. — Left lateral view of the skull of Gyparchus papa; size of life. 

Fig. 117. — Superior aspect of pelvis of Cathartes aura, life size; caudal vertebrae 
still in situ. 



802 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XXII. 

Fig. 118. — Left lateral view of skull of Cathartes aura, size of life; same specimen 
from "which the parts referred to in the other plates were chosen. 
Fig. 119. — Same, viewed from above. 
Fig. 120. — Same, viewed from below. 



804 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XXIII. 

Fig. 121. — Cathartes aura, same as in Fig. 118; right lateral view of sternum ; size 
of life. 

Fig. 122. — Same, seen from below. 

Fig. 123. — Same specimen; dotted outline of furculum, life size, from in front; 
giving the general outline from this aspect of this hone as it occurs in the Cathartidce. 

Fig. 124. — Same specimen (C. aura); right outer aspect of coracoid and scapula; 
life-size. The hone is represented as resting on the plane of the paper. 

Fig. 125. — Same specimen ; right lateral aspect of the furculum, size of life, show- 
ing entrance to pneumatic fossa, and the distribution of the smaller foramina. 



806 GEOLOGICAL SURVEY OF THE TERRITORIES. 



PLATE XXIV. 

Fig. 126. — Left lateral view of the skull of Catliarista atrata ; life size. 

Fig. 127. — Same, from above. 

Fig. 128. — Lower jaw, from above; same specimen. 

Fig. 129. — Same specimen ; tbe skull from below ; life-size. 



i 



